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\' ice-Chairman- M. u. CIIAKRATONC, IONC.YAI 





MRS. M. ISAF^I (II i ARRAR, Secretary 

|'KIN.|ri> A I 1HAI \\A1ANA 1 \NICH ( K l> V >, K\MA I R()A1>, HAN'(.KOK MJ<S HOONPRIN<t F SUW'AN, PRINILR BE 25O5 


1'ldt tor's Note . . .1 

Abbreviations . 11 

Participants ......... in 

Standing Committee Repoits 

IOSBI RG. r.R , Report ofthe Chairman of the Standing Committee on Pacific Botany 1 

WAI KI R, i II., Concerning the Flora of Japan, Fnghsh Translation 4 

nu. SHIN - YiNCj. Status of the Nora of China Project . 5 

\\ AI KI R. i H , Report on the Botanical Section ofthe Program for Scientific Investigation ofthe 

Ryukyu Klands . 6 

si noDiwiim), KDSNOTO, Botany in Indonesia, 1953- 1957 7 

MOORI , i r( \ n , Botany in New /ealand, 1953 - 1957 . 10 

\\AI MR, i n , Report of the Subcommittee on Bibliography 14 

DOIY, MS, Report of the Subcommittee on Algae 17 

STI INI R, MONA i ISA, Report ofthe Subcommittee on Common Names of Pacific Plants IS 

HANK, i. P, II, Report of the Subcommittee on Fthnobolany . . 2! 


ROGI RS. DAVID, Summary of Studies on Mtiiuhot r.senlenta 24 

i AM, H.J., Report of the Subcommittee on Pacific Plant Areas 2N 

SKOI rsiM RG, (".. Report ofthe Subcommittee on Nature Protection . 29 

si i YODIWIRYO, KUSNOTO and DH MY, ANWARI, Problems Confronting Botanical Institutions in 

the Tropics . ......... . ^9 

KOSTI RMANS, A., Problems of Botanical Institutes in the Tropics , . 41 

PtJKSiGi ovi , J.w., A Brief History of Botanic Gardens with Special Reference to Singapore . 44 

BURKILI , n. M., Some Current Problems ofthe Botanic Gardens, Singapore 46 

SUVAFABANDHU, KASiN, Problems Facing the Herbaria in Thailand 52 

VAN STI i MS, c.G.r. j , Interaction and Cooperation between 7 ropical and Temperate Herbaria . 54 

PROWSI , G.A., Problems in a Spceiali/ed Botanical Department . . 56 

VIROI, ROBI RT, The Various Technical Aspects of Botanical Research in New Caledonia 58 

WOMI RSII Y, J.s., Notes on the Techniques Employed for the Collection of Botanical Speci- 
mens in Papua and New Guinea 61 

S\mposium on Vegetation Types of the Pacific 

DANSI KI AU, Pii KKi , Vegetation Study and Recording 64 

HOSOKAVVA, r., OMURA, M. and NISIUHAKA, Y., Grading and Integration of Epiphyte Commu- 
nities ... . ... ... .... 76 

KUC'in [R, A.W , Vegetation Mapping m the Pacific Region S3 

POKU RIS, R., The Problem of the Origin of the "Savannahs" of the Islands of the Pacilic .... 86 

iiiJRi IMANN, J M., The Structure of Some Biocoenoses of New Caledonia . 89 

KICKING, RIIOY w , The hog Belt Rain Forest of the Pacific Northwest (U.S.A.) .... ... 95 

IAII KAWI, MISAO, 'I he Iconography of the Vegetation of the Natural Forest in Japan ... 99 

WANG, rm - vvu. The Development of horest Communities in Eastern Asia . . 103 

IOSBIRG, i R , Dynamics of Atoll Vegetation . . . . 114 

rosBi RG, i .R , Tropical Pacific Grasslands and Savannas .... 118 

KIRA, IAIUO, OGAVVA, iii'SAio and YODA, KYOJI, Some Unsolved Problems m Tropical Forest 

Ecology . . . . . 1 24 

HORIKAWA, Yosiiiwo. The Occurrence of Tropical Plants in the Japanese Archipelago and its 

Phytogeographical Significance . . . .... 135 

i Cii i R, [ RANK i , Unifying Concepts in Vegetation Study as Applied to the Pacilic Basin . 1 36 

VAN STI i NIS, c G G j , The School-Flora as a Medium for Botanical Education m the Tropics 139 

MFIJI R, w.. Botanical Exploration and Education in Present Day Sumatra 141 

HOITIUM, R.I ., Problems of Publicity: with Special Reference to the Need for Popular Books 

dealing with Focal Plants . . 146 

no i Y. MAXWI i i. s., I unctions of the Algae in the Central Pacific 148 

JOHNSON, j. HARI AN, Comparison of the Calcareous Algal Eloras of Recent and Fossil Reefs 156 

IOSBIRG, I.R., Qualitative Description of the Coral Atoll Ecosystem ... 161 

P\LUMBO, RAI PH K, The Relationships between Atolls and Benthic Algae 168 

\\OOD, i j mu.t'soN, The Microbiology of Coral Reefs 171 

c IIAPMAN, v j , 'I he Marine Biogeographical Provinces of the South Pacific 174 

no, PHAM - HOANG, Coastal Marine Plants Around Cauda Harbour (Near Nhatrang) 179 

SCAGI i , ROW KI i ., Benthic Algal Productivity in the North Pacific with Particular Reference 

to the Coast of British Columbia 181 

RYrm:R, JOHN H., On the Efficiency of Primary Production in the Oceans ... 188 

sii v\, PAUL c.. Comparison of Algal Floristic Patterns in the Pacific with those in the Atlan- 
tic and Indian Oceans, with Special Reference to Codium .... 201 

PROWSI , G.A., Distributional Relationships of Malayan Freshwater Algae 217 

in DMANN, Ji AN, The Rhodophyta Order Acrochactiales and its Classification ... . ... 219 

IOKIDA, JUN and YAIHJ, IIIROSHI, Some Observations on Lammaria Gametophytes and Sporo- 

phytcs 222 

PAPINIUSS, GLORGI K, Clearing Old Trails in Systematic Phycology . 229 

YI i ASQIII /, GRi.GORio i.. On the State of Phycological Knowledge in the Philippines 234 

SCAGLI , ROBi RT p., Phylogcnctic Relationships of Certain Dorsivcntral Rhodomelaceae .... 239 

Symposium on I'thnohotain of 1 lunland and Contiguous Countries 

BANK, IHIODORI P . II, Opening Remarks on Hthnohotany and Ecology . 244 

JONIS, \OIMY ii.. The Nature and Status of Fthnobotany . . ... . 246 

SMITINAND, ii M, Materials Used for Thatching m Thailand . ... 248 

SAMAPUDDIII, k., Some hood Plants in the ForesN of 'I hailand . . . 250 

SMIIINAND, TI M, Note Made from Local Knowledge of the Use of Poisonous Plants by the Thai 

People . . . . 262 

PHDMXUSRI, pRASimu, Tea in Thailand . . . 264 

IUGBY, nisi. The Manufacture of Sugar from the Sugar Palm in Upper Mandailing, 

Sumatra . . .... 266 

BART i ITT, IIAKI i Y HARRIS, Possible Separate Origin and Evolution of the Ladang and Sawah 

Types of Tropical Agriculture 270 

BARii.Lir, HARI i/v HARRIS, Some Words Used in Connection with Primitive Agriculture in 

Southeast Asia . ... 274 

WIATHFRWAX, PAUi , Ethno-Ecological Relationships between the Mai/e Plant and Man in 

Western Ancient America 276 

BANK, IHLODORF P., II, Elhiiobotany of Northern Peoples and the Problem of Cultural Drift . . 279 

BANK, THLODORE p., II, Medicinal Plant Lore of the Aleut 281 

STHINL R, MONA i ISA, The Problem of Vernacular Names of Plants in the Pacific and its Solution 285 

HU, smu - YING, An Enumeration of the Food Plants of China with Vernacular Names 289 

TUYAMA, TAKASI, The Problem on the Vernacular Plant Names in Japanese 291 

PURSLGLOVF, J.w., Common Vernacular Names of Plants of the Pacific Basin 293 

BARRAU, JACQUES, Notes on the Significance of Some Vernacular Names of Food Plants in 

the South Pacific Islands 296 

CONKLIN, HAROLD c., Ethnobotanical Problems in the Comparative Study of Folk Taxonomy. . 299 

AUSTERLITZ, ROBERT, A Linguistic Approach to the Ethnobotany of South-Sahalin 302 

DWYHR, R.E.P., Vernacular or Common Names of Plants 304 

JOHN, HAROLD ST., Origin of the Sustenance Plants of Polynesia, and Linguistic Evidence for 

the Migration Route of the Polynesians into the Pacific (Abstract) 308 

BARRAU, JACQUES, Edible Yams of the South Sea Islands, Species Present, Vernacular Names 

and Distribution 309 

Symposium on Vernacular Names of Plants 

COLE, KATiiihLN, Acelo-Carmine Stain in the Cytogemitical Investigation of Some Marine 

Algae of the Pacific Coast 313 

ELLIOTT, JACK c., The 1956 Typhoon Season on Okinawa (Abstract) 316 

TIENG, TRAN-NGOC, The Evolution of the Germinative Power of Rice Seeds 318 

DE JONGI , P., Stimulation of Yield in Hevea Brasiliensis 320 

MENNINGER, EDWIN A., Flowering Trees that Fail to Flower in Alien Lands 325 


At the meeting of the Committee on Publications of the Ninth Pacific Science Congress, it was 
agreed that the increasing number of communications and papers presented at each Congress has 
become a very difficult problem for the Publication Committee and the editorial staff to cope with and 
that too much time is required to complete the publication of the Proceedings; therefore, it was recom- 
mended that the following principles governing publication be followed: 

a. That invited contributions to the scheduled symposium be published in full; 

b. That reports of the Standing Committees be published in full; 

c. That other papers submitted to the Congress during its sessions be published in abstract only, 
the abstract not to exceed 500 words; 

d. That papers which, though listed on the program or included in the pre-Congress abstracts 
published in advance but not actually submitted to the Congress at its sessions, should be 
disregarded ; 

e. That authors be asked to indicate by a definite and early date 1 whether they prefer to publish 
their papers in sources other than the Congress Proceedings; that if this is done, the Congress 
should be acknowledged; 

f. That all proof reading be the responsibility of the editorial committee, and that this committee 
shall consider the manuscripts in their hands by a definite date as final; 2 

g. That authors be held responsible for submitting their material in good English; 

h. That on matters arising during the course of publication and not specifically covered in the 
statement of policy the editorial committee is empowered to act. 

In accordance with the resolutions of the Committee, the editorial board has edited the reports and 
manuscripts where necessary to bring uniformity and consistency to the format. Typographical and 
grammatical errors as well as errors in phraseology, spelling, or technical terms have been corrected, 
wherever possible, but in cases where the exact meaning of the original copy was not clear, the text has 
been left as submitted by the autor. 

In order to reduce the cost and bulk of the publication, appendices, illustrations, and exhibits when- 
ever considered not vital to the text have been eliminated. 

Tf an author requested to publish elsewhere, his paper has been mentioned in the footnote under 
the respective titles, but if an author who presented a paper at the Congress failed to submit his manu- 
script either iafull or in abstract, his paper and the discussions thereon have been eliminated entirely. 

It was also decided that, in order to complete the publication of the Proceedings as soon as possible, 
each division be published in a separate volume. Short volumes or the ones that do not require too much 
editorial work will be released first. Therefore, among the twenty volumes planned, any volume may 
appear first. They will not appear in consecutive order. 

The editorial board wishes to thank all authors who were prompt in submitting their revised manu- 
scripts in good form and, in particular, members of the Standing and Organizing Committees, too 
numerous to be named, who have helped in collecting the manuscripts pertaining to their respective 

The Board wishes in particular to thank Dr. F. Raymond Fosberg for going over and correcting 
the Special Symposium on Climate, Vegetation, and Rational Land Utilization in the Humid Tropics 
under UNESCO; 

Mr. Saman Buravas of the Royal Mines Department for helping by redrawing charts and maps 
in order that they might reproduce clearly when printed; 

Mr. J. Alan Tubb of the FAO Regional Office, for his assistance in going over and clarifying some 
of the papers in the Fisheries and Oceanography volumes and in translating some of the French papers; 

Dr. Pradisth Cheosakul of the Department of Science for editing the Chemistry in the Development 
of Natural Resources volume; 

Last but not least, the Board wishes to thank the Thai Watana Panich Press for their cooperative ef- 
forts, far beyond the requirement of the contract, in devoting all their resources to printing these volumes. 

1 January 1, 1958, in the case of the Ninth Congress. 

2 March 1, 1958, in the case of the Ninth Congress. 





























Asia-Pacific Forestry Commission 

Civil Air Administration 

Commonwealth Scientific and Industrial Research Organization (Australia) 

Economic Commission for Asia and the Far East 

Equatorial Pacific (occanographic survey) 

Food and Agriculture Organization 

International Advisory Committee on Marine Sciences 

International Cooperation Administration 

International Civil Aviation Organization 

International Council of Scientific Unions 

International Geophysical Year 

Indo-Pacific Fishery Commission 

International Rice Commission (FAO) 

Joint Commission on Rural Reconstruction (Taiwan, China) 

North Pacific (occanographic survey) 

Philippine Council for United States Aid 

Pan-Indian Ocean Scientific Association 

South-East Asia Treaty Organization 

South Pacific Commission 

United Nations 

United Nations Educational, Scientific and Cultural Organization 

United Nations International Children's Emergency Fund 

United States Department of Agriculture 

United States Information Service 

United States of America Operations Mission 

World Health Organization 

World Meteorology Organization 


ASHTON, PETER SHAW, Botanist, State Forest Office, Brunei. 

AUBERT DE LA RUE, EDGAR, Geologist, National Centre for Scientific Research, 18 rue Ribera, Paris 
XVI", France. 

BALASIRI, DAROON, Instructor, School of Pharmacy, University of Medical Science, Bangkok, 

BANK, THEODORE FOUL, ii, Executive Director, Institute for Regional Exploration, University Station, 
Box 2143, Ann Arbor, Michigan, U.S.A. 

BARTLETT, STEPHEN, Head of F&M Department, National Institute for Research in Dairying, Univer- 
sity of Reading, Reading, England. 

BELARDO, LUZ ouvEROS, Dean, College of Pharmacy, The Philippine Women's University, Manila, 

BOONJAVATANA, CHAROEN, Supervisory Unit, Department of Teacher Training, Ministry of Educa- 
tion, Bangkok, Thailand. 

BRYNE, BRIAN, Teacher, Mt. Hermon, Darjeeling, India. 

BURKILL, HUMPHREY MORRISON, Director, Botanic Garden, Singapore. 

CALDWELL, ARTHUR FRANCIS, Department of Pharmacy, Faculty of Medicine, University of Malaya, 

CAPELI , ARTHUR, Reader in Oceanic Linguistics, University of Sydney, Sydney, N.S.W., Australia. 

CHANTARASRIKUL, ANONG, Bangkhen Experimental Station, Department of Agriculture, Ministry of 
Agriculture, Bangkok, Thailand. 

CHUMDFRMBHADEJSUK, SHALAM, Srisuriyothai School, Ministry of Education, Bangkok, Thailand. 
CROCKER, ROBERT LANGDON, Professor of Botany, University of Sydney, Sydney, N.S.W., Australia. 
DASANANDA, SALA, Deputy Director-General, Rice Department, Ministry of Agriculture, Bangkok, 


DOTY, MAXWELL s., Professor of Botany, University of Hawaii, Honolulu 14, Hawaii. 
DOUGLAS, BRYCE, Senior Scientist, Smith Kline and French Laboratories, Philadelphia, c/o Chemistry 

Department, University of Malaya, Singapore 10. 
EGLKR, FRANK EDWIN, Aton Forest, Norfork, Connecticut, U.S.A. 
FELDMANN, JEAN, Professeur & la Faculte de Sciences de Paris, Laboratoire de Biologic Vegetale 

Marine, Faculte dc Sciences de Paris, 195 rue St. Jacques, Paris 5 e , France. 

FOSBERG, F. RAYMOND, Botanist, U.S. Geological Research Council, Washington 25, D.C., U.S.A. 
GAUSSEN, HENRI MARCEL, Professor, University of Toulouse, rue Raymond JV 21, Toulouse, France. 
HARRISSON, TOM H., Curator, Sarawak Museum, Brunei. 
HEIM, ROGFR, Director of the National Museum of Natural Sciences, Museum National D'Histoire 

Naturelle, 57 rue Cuvier, Paris 5 e , France. 

HO, PHAM HOANG, Charg de covas a la Faculte, Universite de Saigon, 12 Hong Thap Tei, Saigon, 

HOERNER, GODFREY RICHARD, Chief University Advisor, Kasetsart University/Oregon State College 

ICA Contract, Bangkhen, Bangkok, Thailand. 

HORMCHONG, TWFE, Instructor, College of Education, Ministry of Education, Bangsaen, Thailand. 
HU, SHIU-YING, Botanist, Arnold Arboretum, Harvard University, 22 Divinity Ave., Cambridge, 

Mass., U.S.A. 
HURLIMANN, JOHANN H., Botanist, Ciba J.A., Friendensgasse 51, Basle, Switzerland. 

JALAVICHARANA, KAHN, Chief, Plant Industry Division, Department of Agriculture, Bangkhen, 
Bangkok, Thailand. 

JIRAWONGSE, VICHIARA A., Lecturer, Department of Pharmacy, Chulalongkorn University, Bangkok, 

* Initials or names in italics represent Thai titles. 


JOHNSON, J. HARLAN, Professor of Geology, Colorado School of Mines, Golden, Colorado, U.S.A. 
KASHEMSANTA, M.C. LAK, Director-General, Department of Agriculture, Ministry of Agriculture, 
Bangkok, Thailand. 

KETUSINH, OUAY, Professor, Department of Physiology, Siriraj Hospital, Dhonburi, Thailand. 

KUCHLER, A.W., Professor of Geography, University of Kansas, Lawrence, Kansas, U.S.A. 

LOETSCH, FRITZ, Professor, Hamburg University, Bundes forschungsanstalt Forstwirtschaft, Reinbek 
bei, Hamburg, Germany. 

MCHALE, THOMAS RiLEY, Economist, 24 Blackstone Blud., Providence, R.I., U.S.A. 

MEKAYONK, VICHIT, Suan Sunantha College, Bangkok, Thailand. 

NA-THALANG, SOMBOON, Director, Rubber Division, Department of Agriculture, Ministry of Agri- 
culture, Bangkok, Thailand. 

OGAWA, HUSATO, Researcher, Laboratory of Plant Ecology, Institute of Polytechnics, Osaka City 
University, Manami-Ogimachi, Kitaku, Osaka, Japan. 

PAPENFUSS, GEORGE F., Professor, Department of Botany, University of California, Berkeley 4, Cali- 
fornia, U.S.A. 

PAVANARIT, SRIROT, College of Education, Department of Teacher Training, Ministry of Education, 
Bangkok, Thailand. 

PHUMXUSRI, PRASIDHI, Agriculturist, Mae Malai Farm, Mae Rim, Chiang Mai, Thailand. 

PICHAISIRI, SOMSUGDI, Instructor, Department of Teacher Training, Ministry of Education, Bangkok, 

PROWSE, GERALD ALBERT, Botanist and Algologist, Fish Culture Research Station, Batu Berendam, 
Malacca, Federation of Malaya. 

RAYMOND, MARCEL L., Taxonomist, Montreal Botanical Garden, 4101 East, Sherbrooke Street, Mon- 
treal, Canada. 

RICHARDS, PAUL WESTMACOTT, Professor of Botany, University College of North Wales, Bangor, 
Great Britain. 

RUHLE, GEORGE CORNELIUS, Park Naturalist, Hawaii National Park, Hawaii, U.S.A. 

RYTHER, JOHN HOOD, Marine Biologist, Woods Hole Oceanographic Institution, Woods Hole, Mass., 

SACHET, MARIE-HELENE, Bibliographer, Pacific National Research Council, Washington 25, D.C., 

SAMAPUDDIH, KRiT, Royal Forest Department, Bangkok, Thailand. 

SANSUKH, SOMSAK, Teacher, College of Education, Department of Teacher Training, Ministry of 
Education, Bangkok, Thailand. 

SANTOS, JOSE VERA, Head, Department of Botany, University of the Philippines, Quezon City, 

SCAGEL, ROBERT FRANCIS, Associate Professor, Department of Biology and Botany, and Institute of 
Oceanography, University of British Columbia, Vancouver 8, B.C., Canada. 

SEIDENFADEN, GUNNAR, Danish Minister, Danish Legation, Bangkok, Thailand. 

SETYODIWIRYO, KUSNOTO, Director, Botanic Gardens of Indonesia, Djl. Bubulak 25, Bogor, Indonesia. 

SILVA, PAUL CLAUDE, Associate Professor, Department of Botany, University of Illinois, Urbana, 
Illinois, U.S.A. 

SMITINAND, TEM, Chief, Section of Botany and Zoology, Forest Products Research Division, Royal 
Forest Department, Bangkok, Thailand. 

STEINER, MONA LISA, Chairman of sub-committee on Vernacular Names of Plants, 2833 Park Avenue, 
Pasay City, Philippines. 

SUVAGONDHA, CHAMLONG, Professor and Dean, School of Pharmacy, Bangkok, Thailand. 

SUVARNASUDDHI, KHio, Deputy Director-General, Royal Forest Department, Bangkok, Thailand. 

SUVATABANDHU, KASIN, Department of Agriculture, Bangkok, Thailand. 

THIRAWAT, SUKHUM, Chief, Division of Forest Control, Royal Forest Department, Bangkok, 


THONGUMPHAI, KHAJORN, Teacher, Chachoengsao Teacher Training College, Chachoengsao, 


TRINANDWAN, SA-KORN, Teacher, College of Education, Ministry of Education, Bangkok, Thailand. 
TROLL, CARL TH., Professor, University of Bonn, Universitat, Geographisches Institut, Bonn, Germany. 
TUYAMA, TAKASI, Professor, Laboratory of Botany, Faculty of Science, Ochanomizu University, 

Otsuka, Bunkyoku, Tokyo, Japan. 

VALLARTA, MARIA T., Head, Department of Botany, The Philippine Women's University, Manila, 

VAN STEENIS, c.G.G.J., Professor, Flora Malesiana Foundation, Marelaan 55, Oegstgeest, Netherlands. 

VESTAL, EDGAR F., T.A. Plant Protection Advisor, USOM, Bangkok, Thailand. 

WALKER, EGBERT H., Department of Botany, Smithsonian Institution, Washington 25, D.C., U.S.A. 

WEATHERWAX, PAUL, Visiting Professor, College of Education/Indiana University ICA Contract, 
Bangkok, Thailand. 

WYCHERLEY, PAUL RENODEN, Botanist, Rubber Research Institute of Malaya, P.O. Box 150, Kuala 
Lumpur, Malaya. 

YEN, DOUGLAS ERNEST, Scientific Officer, Vegetable Station, D.S.I. R., Robinson Road, Otahuhu, 
New Zealand. 

YODA, KYOJI, Researcher, Laboratory of Plant Ecology, Institute of Polytechnics, Osaka City Uni- 
versity, Minami-Ogimachi, Kitaku, Osaka, Japan. 


Standing Committee Chairman: F. R. FOSBERG 
Organizing Committee Chairman: M.C LAKSHANAKARA KASHEMSANTA 

Standing Committee Reports 




Chairman, Pacific Vegetation Project, Washington, D.C., U.S.A. 

The Standing Committee on Pacific Botany, 
established under this title by the Eighth Pacific 
Science Congress, succeeding the former Standing 
Committee on Pacific Plant Areas, includes the 
following members (arranged alphabetically): 

Theodore P. Bank II (U.S.A.) (Chairman, 
Subcommittee on Pacific Ethnobotany). 

Nancy T. Burbidge (Australia). 

Pierre Dansereau (Canada) (Chairman, Sub- 
committee on Pacific Plant Ecology). 

Maxwell S. Doty (Hawaii) (Chairman, Sub- 
committee on Pacific Algae). 

Ramon Ferreyra (Peru). 

F. Raymond Fosberg (U.S.A.) (Chairman, 
Standing Committee). 

Hiroshi Kara (Japan). 

Ir. Kusnoto Setyodiwiryo (Indonesia). 

H. J. Lam (Netherlands) (Chairman, Subcom- 
mittee on Pacific Plant Areas). 

E. Looho (Indonesia) (Chairman, Subcommit- 
tee on Medicinal Plants). 

Lucy B. Moore (New Zealand) (replacing the 
late W.R.B. Oliver). 

Eduardo Quisumbing (Philippines). 

Carl Skottsberg (Sweden) (Chairman, Subcom- 
mittee on Nature Protection). 

Mona Lisa Steiner (Philippines) (Chairman of 
Subcommittee on Common Names of Pacific 

Kasin Suvatabandhu (Thailand) (Secretary, 
Standing Committee). 

Egbert H. Walker (U.S.A.) (Chairman, Subcom- 
mittee on Botanical Bibliography). 

John Wyatt-Smith (Malaya). 

The committee has functioned principally 
through its eight subcommittees. The reports of 
these subcommittees (which follow) speak for 

themselves, and material contained therein will 
not be repeated here. Their accomplishments 
are amply sufficient to justify continuation of this 
method of functioning of the Standing Com- 
mittee. It is therefore recommended that the 
Botany Division of the Ninth Congress consider 
very carefully the lines of effort that it wants the 
next Standing Committee on Pacific Botany to 
emphasize and that it recommends for each such 
line or field the establishment of a special sub- 
committee, as was done in 1953. 

The present Standing Committee is composed 
of the chairmen of the subcommittees and suffi- 
cient other members to give it wide geographical 
representation. No members were selected from 
countries that were represented already by sub- 
committee chairmen. When Professor Kusnoto 
had to resign the chairmanship of his Subcom- 
mittee on Medicinal Plants because of pressure 
of other duties, on his recommendation Dr. 
Looho was appointed to replace him, but he 
continued as a Committee member. We deeply 
regret the loss by death of one of our members, 
Dr. Oliver, chairman of the predecessor of this 
committee. Miss Moore was asked to replace 

The principal efforts of the Chairman were 
to assist the Organizing Committee of the Ninth 
Congress with the planning of the botanical 
program and to compile a list of Pacific botanists. 

To accomplish the first aim, a visit was made 
to Bangkok in April, 1956, when the program was 
thoroughly discussed and names were suggested 
of people who might agree to organize symposia 
and to be chairmen of sessions. Most of these 
people have responded to the invitations of the 
Organizing Committee, resulting in the rather 



ambitious program of the Botany Division, 
and, incidentally, in the excellent representation 
of Pacific botanists at the Congress. Where 
possible, the activities of the subcommittees have 
been tied into the program. If certain fields have 
been neglected, it was because the program could 
scarcely have been made longer. Cooperation 
with other divisions was recommended where it 
seemed appropriate. 

The second of the tasks was more difficult, 
and the product is admittedly imperfect. A 
request was sent to all Standing Committee 
members and a few others for lists of names from 
their several geographic areas. This resulted 
in a few lists. To these were added names known 
to the Chairman, encountered in the literature, 
or found in published address lists. The resulting 
list was duplicated and circulated to a geograph- 
ically representative group of botanists for 
criticism, additions, and corrections. The result 
was gratifying and enabled us to get together the 
list that is before you and which will be mailed 
to all those appearing in it. It is hoped that it 
will contribute substantially to the primary aim 
of the Standing Committee, to stimulate com- 
munication and cooperation between the botanists 
interested in the Pacific Basin. 

A word is in order about the scope and limita- 
tions of this list. It was arbitrarily limited to 
those branches of botany that have a definite 
regional aspect, and that are not properly included 
in agriculture or other strictly practical fields 
which may have the resources to compile lists 
that will better serve their purposes. Thus many 
names were excluded in such fields as plant phy- 
siology, genetics, plant chemistry, cytology, 
agronomy, etc. Reliance usually had to be 
placed on printed sources for such information. 
Undoubtedly serious errors were made. We 
humbly ask forgiveness and hope that a future list 
may rectify all errors and omissions. 

We take great pleasure in acknowledging our 
gratitude to the Pacific Science Council Secreta- 
riat and the National Science Foundation for 
financial help that made the production of this 
list possible. Copies will be distributed to all 
people whose names appear on the list and to 
libraries of institutions known to be interested 
in Pacific Botany. Persons or institutions not 
listed or which do not receive copies may receive 
them by requesting them, so long as the copies 

We wish also to express our gratitude to the 
Pacific Science Board of the National Research 
Council for continuously making facilities and 


space available for the functioning of the Stand- 
ing Committee, and for help in numerous ways. 

It is customary for the reports of standing 
committees to include summaries of work in the 
field accomplished or undertaken during the 
period of the Committee's activity. We have 
asked Committee members for such information 
on their areas. A few have responded, and their 
contributions are briefly summarized here, along 
with some other widely known current activity. 
This is only a sampling of the field, and details 
and most names are purposely omitted, as it is 
impossible to list all of them. Several of the 
reports submitted by Committee members seem 
sufficiently comprehensive that it is desirable to 
append them in full to this report, in which cases 
little in the way of summary is included. 

By far the most comprehensive and widespread 
program in the Pacific Area is that of the Flora 
Malesiana, whose able editor is with us and may 
care to give us current news of it. Under its 
auspices have appeared many preliminary re- 
visions of groups as well as a number of the 
final treatments for the Flora. The Flora male- 
siana Bulletin continues to be the world's most 
informative botanical news organ, as well as 
containing much valuable bibliographic informa- 

Revision of Cheeseman's Manual of the Flora 
of New Zealand is complete, and the book is in 
press. For further information on New Zealand, 
see Miss Moore's report. 

The C.S.I.R.O. in Australia is carrying out 
extensive botanical and ecological survey work in 
the Northern Territory and in New Guinea, 
and botanists in other parts of Australia continue 
to be very active. It should be specially noted 
that the botanical exploration of the subantarctic 
Macquarie Island, recommended by the Second 
Pan-Pacific Science Congress, has been carried 
out by a group from the C.S.I.R.O. 

Along the Pacific side of South America, work 
on the flora and vegetation of Peru, centered at 
the Universidad de San Marcos, Lima, and that 
on Colombia, centered at Bogota, Medellin, 
and especially at the U.S. National Herbarium, 
is especially active and productive. The Flora of 
Panama is being continued by the Missouri 
Botanical Garden. 

In the U.S., work on a manual of California 
plants is going on at the Rancho Santa Ana 
Botanical Garden and New York Botanical 
Garden. At the several campuses of the Univer- 
sity of California, Jepson's Flora of California 
is drawing toward completion, and much work in 



systematic, biosystematic, and ecological fields 
is being done. Pomona College and affiliated 
institutions also form an active center of taxo- 
nomic and ecological work, especially on plants 
of arid southwest North America. Work on the 
vegetation of Alaska has been carried out under 
various auspices, especially in connection with 
work on permafrost by several military sponsored 
agencies and the U.S. Geological Survey. 

In Japan there has been much activity in many 
institutions, principally on the vegetation and 
flora of Japan. Especially valuable as a basis for 
all other such work are Hara's bibliographic 
checklist of the flora of Japan and an English 
language revision of Ohwi's Flora of Japan, the 
latter in cooperation with several U.S. Institu- 

We have no news of the status of recent 
botanical work in mainland China, but the Flora 
of China Project of the Arnold Arboretum, 
U.S.A., is being continued under the direction of 
Dr. Shin-ying Hu. (See accompanying report by 
Dr. Hu.) 

For a report of progress on a flora of the 
Ryukyu Islands see the accompanying report by 
E.H. Walker. 

In Thailand aquatic and economic plants are 
under study by our Thai colleagues, and a Danish 
expedition has been studying and collecting the 

At the University of Hawaii is a project on 
Pacific Pandanus, as well as active work on algal 
ecology and ocean productivity. 

The Pacific Vegetation Project has continued 
its investigations on the vegetation and flora of 
Micronesia, as well as some other Pacific areas 
and on coral atolls generally, sponsored by the 
U.S. Geological Survey. A by-product of this 
is the volume, Island Bibliographies, published 
by the Pacific Science Board. 

On other insular areas, work on the flora of 
New Caledonia has continued at Zurich and by 
R. Virot at Paris, on that of Fiji at the U.S. 
National Herbarium and at Suva, and on the 
food plants of the Pacific at the University of 
Marseilles and the South Pacific Commission. 
Finally, we must mention the completion and 
publication of two magnificent volumes on the 
vegetation and the origin and affinities of the flora 
of Juan Fernandez by our eminent colleague, Carl 
Skottsberg of Gothenburg, Sweden. 

Mention should be made of the status of the 
volume on the Vegetation Provinces of the 
Pacific Basin, based on the symposium at the 
Eighth Pacific Science Congress. Manuscripts 
are in hand for the greater part of the chapters, 
but about a third are still not available. They 
are mostly reported in various stages of comple- 
tion. It is hoped that this work can be promptly 
brought to a conclusion, as all indications are 
that the volume is much needed and would imme- 
diately be widely used. Experience with this 
volume tends to discourage attempts at coopera- 
tive works of this sort, as it seems to be necessary 
to spend time out of all proportion merely to 
urge unenthusiastic collaborators to complete 
their promised contributions. 




Smithsonian Institution, Washington, D.C., U.S.A. 

An American serviceman, Leopold Charette, 
initiated the translation into English by J. Ohwi 
of his Flora of Japan (in Japanese, 1953). Fred 
Meyer, formerly at the Missouri Botanical 
Garden, and E.H. Walker, Smithsonian Institu- 
tion, are editors, and the project is currently 
sponsored by the Missouri Botanical Garden. 
Funds have been provided by the National 

Science Foundation, U.S.A. Dr. Ohwi recently 
submitted a treatment of the Pteriodophyta, a 
group not included in the original flora, in 
English. With the permission of the editors, he 
then translated this into Japanese for publica- 
tion in Japan. This came from the press in 
November, 1957. (A copy was circulated through 
the audience.) 





Arnold Arboretum, Jamaica Plain, Massachusetts, U.S.A. 


This project was initiated in 1953 as a coopera- 
tive project between A Continental Development 
Foundation and the Arnold Arboretum of 
Harvard University. Over thirty thousand 
dollars has been invested in it. 


1. A complete index to the species of the 
flowering plants has been prepared. This is in 
card catalogue form and it consists of approxi- 
mately 100 thousand cards each covering names, 
synonyms, important literature references, au- 
thentic collections, and the distribution. 

2. The preparation of a bibliographical enu- 
meration of the plants of China has commenced. 
The treatment of the Compositae is now complete. 

The manuscript consists of about 1,500 typed 
sheets, double spaced. That of the Orchidaceae 
is almost done. 


1. A model of the flora, a treatment of the 
Malvaceae, was published in 1955 and distributed 
in 1956. It shows what can be done. 

2. Manuscript for the next issue covering 
Palmae, Araceae, Ericauloneceae, Conmelina- 
ceae, and Panderderaceae is at hand. 


The supporting foundation can no longer 
provide financial support to the Project. The 
undertaking itself is a tremendous task. Now 
moral and financial support are required for its 




Smithsonian Institution, Washington, D.C., U.S.A. 

The SIRI program of the Pacific Science Board 
and the U.S. Army was designed to provide 
basic scientific information for the development 
of the southern Ryukyu Islands. As botanist on 
this program, I spent the three summer months 
of 1951 in Okinawa and the Southern Ryukyus 
mainly collecting herbarium material for use in 
eventually preparing a Flora of the area. 

In 1952, the U.S. Civil Administration and the 
Government of the Ryukyus mimeographed an 
issue of 200 copies of a Flora of Okinawa, 
originally prepared by the Okinawans, Sonohara, 
Tawada, and Amano, and edited by me. In 1954, 
they issued my book of some 300 pages Important 
Trees of the Ryukyu Islands, dealing with 209 
species, each illustrated, the text in both Japanese 
and English in parallel columns. In 1953, I 
attended the Eighth Pacific Science Congress as 
delegate from this program and contributed 
two papers dealing with Okinawa and the 
Southern Ryukyus, and on my return made 
further collections in Okinawa. With assistance 
from this program, a herbarium was established 
at the University of the Ryukyus in 1955 and 
encouragement given to the training in America 
of an Okinawan botanist to become its curator. 

In 1954, a new Flora of Okinawa and the 
Southern Ryukyus was started. It is based upon 
all available specimens, which have now grown 
to occupy some ten cases in the U.S. National 
Herbarium in the Smithsonian Institution, and 
on critical studies by many contributors. As 
now being prepared it will contain keys, brief 
characterizations, and critical notes. The thor- 
oughness of this work was greatly increased by 
the opportunity to select and borrow critical 
herbarium specimens from Japanese and Taiwan 
herbaria and to make collections in Okinawa 
of palms and bamboos while enroute to this 

Besides the material results of this program, 
so useful in the development of the Ryukyus and 
of our scientific knowledge of this area, such as 
the publications and critically studied scientific 
specimens, certain intangibles may be noted. 
Especially significant is an increased under- 
standing, goodwill, and cooperation between the 
botanists and institutions of Japan, Okinawa, 
Taiwan, America and some European countries. 
Surely the investment in this program is bringing 
rewarding returns. 




Director, Botanic Gardens of Indonesia, Bogor* Indonesia. 


This is a brief report of botanical activities in 
Indonesia during 1953-1957. 

The staff of the Treub Laboratory was engaged 
in conducting the following scientific investiga- 


(a) Research with growth hormones, isolation 
of growth promoting compounds of a fern, 
application of a new Ageratum test, applica- 
tion of commercial growth hormones for the 
rooting of cuttings of cultivated plants. 

(b) Investigations on the interaction between 
epiphytes, micro-organism, and host-plants 
in order to clarify a supposed symbiotic ef- 
fect. Microscopy of the lutoids in Hevea 

(c) Cytological investigations of the Amorpho- 
phallus and Corypha. Description of the 
meiotic divisions, the explanation of a new 
phenomenon found in the meiotic prophase 
in the genus Corypha, in respect of the 
function of the nucleolus. Study on the 
pollen grain mitosis of Raffles ia arnoldi, 
and the establishment of the number of 
chromosomes. Study on the sterility of 
Monodora myristica. Allium spp. were 
used to test the effect of the digitonin on 
the nuclear division. 

(d) Genetical investigations with Digitalis spp. 
carrying out interspecific crossing between 
D. lanata and D. lutea. Observation of 
different Digitalis spp. cultivated on higher 

(e) Anatomical investigations on the leaf and 
root structure of Fagraea borneensis and 
Smilax sp. 


(a) Investigations of the chemical composition 
of latex in the family of Apocynaceae. 

(b) Determination of the rubber hydrocarbon 
constituent of Balanophora sp. 

(c) Investigations on the cardiac glycosides of 
Theyetia peruviana, Antiaris toxicaria and 
various other plants of the families of Ascle- 
piadaceae, Apocynaceae, Moraceae. Test 

for the presence of alkaloids and saponins 
in plants collected in Indonesia. 


(a) Isolation of pure cultures of Pseudomonas 
sp. collected from diseased Hevea and 
Digitalis plants. 

(b) Investigation of Xanthomonas sp. isolated 
from infected rice plants. 

(c) Obtaining of new material for the collection 
of fungi and bacteria maintained in the 
laboratory for scientific and industrial pur- 
poses. Exchange of samples with institutions 

Lectures. Delivering a course on the cultivation 
methods of orchids in 1956. Lectures on Cyto- 
logy and Genetics at the Faculty of Agriculture 
of the University of Indonesia. The same lec- 
tures, and also on Biochemistry, were given at the 
Biological Academy, established by the Ministry 
of Agriculture in October, 1955, in Bogor. 

Excursions. An excursion was carried out to the 
islands of Timor, Roti, etc. in order to collect 
medicinal plants; another excursion in Java, 
Celebes and in the Lesser Sunda Islands to collect 
cytological material of monocarpic palms. 

Guest research workers in the Treub Laboratory. 
Prof. Dr. Koernicke, Germany, Prof. Dr. F. 
Fagerlind and G. Wibom, Sweden, Prof. Dr. P. 
Buchner, Italy, and Dr. P. Surany, U.S.A., have 
visited and worked for varying periods. Dr. 
H.P. Bottelier, Holland, worked in the Treub 
Laboratory on a Fellowship for six months. 

Exhibition. The Treub Laboratory has partici- 
pated in the Agricultural and Horticultural 
Exhibition at Pasar Minggu near Djakarta in 
1954, with a show material on the effect of the 
growth hormones and the symbiosis between 
epiphytes, microorganism, and host plants. 


In 1953, a combined expedition of Dr. A. 
Kostermans of the Forestry Service and Dr. W. 
Meyer of the Herbarium went to the island of 
Nunukan in North East Kalimantan (Borneo) 



and the adjacent mainland. The collections 
amounted to 1,500 numbers of Phanerogams and 
600 Cryptogams. Two other members of the 
staff, Mr. Groenhart and Mr. Van Borssum 
Waalkes, made a trip to West-Central Sumatra 
and the Mentawei Islands. The Herbarium 
received 4,630 plants in exchange, whereas more 
than 14,000 specimens were sent abroad as 
duplicates and more than 8,000 sheets were 
loaned for study purpose. One part of Volume 2 
of Reinwardtia appeared with monographies and 
papers on Phanerogams. 

In 1954, a guest worker, Dr. Alston from the 
British Museum in London, made collecting trips, 
sponsored by the Herbarium, to South Borneo, 
Sumatra, North Celebes, Ternate, and Ambon. 
A trip by Mr. A. Groenhart was made to Banten 
in West Java. The collections of the expedition 
of Dr. Kostcrmans of the Forestry Service to 
Central Kalimantan, amounting to 1,500 field 
numbers, were ceded to the Herbarium. More 
than 3,000 duplicates were received from abroad, 
in exchange for almost 12,000 duplicates, sent 
abroad to twenty Institutes. One big volume 
appeared of Reinwardtia containing numerous 
articles on Phanerogams and Cryptogams. 

In 1955, an expedition of Dr. A. Kostermans 
went to Central Kalimantan for three and one- 
half months. The collections, consisting of about 
one thousand field numbers, are incorporated 
in the Herbarium collections. Furthermore, an 
expedition of Dr. Kostermans to Central Suma- 
tra for the purpose of collecting timber for re- 
search in cellulose for rayon, brought back more 
than 600 field numbers for the Herbarium. Dr. 
Kostermans was accompanied during the first 
part of this trip by Mr. Anwari Dilmy, Keeper 
of the Herbarium. The two gentlemen accom- 
panied Prof. Kusnoto, the Director of the Botanic 
Gardens to Padang Tinggi, West Coast of 
Sumatra, for the official opening ceremonies of 
the Branch Garden there. Plans were outlined 
as to how to investigate the extensive area of 
virgin forest surrounding this new Garden. 
Almost 20,000 specimens have been sent abroad, 
on loan and as gifts, to thirty Institutes, whereas 
3,000 specimens apart from the Kostermans 
collections were received in exchange. One 
back-volume, containing the plates of Ebenaceae 
of Bakhuizen's monography appeared, and one 
part of Reinwardtia, containing numerous taxo- 
nomical articles. The Herbarium building un- 
derwent big repairs and improvements and is 
now in a good condition. 

In 1956, Mr. Van Borssum Waalkes made an 
expedition to Timor. Mr. Jacobs made a 


combined trip with Dr. Meyer from the Agricul- 
tural Faculty at Pajakumbuh to Mount Korintji 
in Central Sumatra, whereas one guest-worker, 
Mr. Forman from Kew, made trips to East Java 
with Mr. Dilmy, and to Sumatra. Dr. Koster- 
mans, who made an expedition to the mountain 
region in Central Kalimantan, brought back 
about 800 field numbers and living plants of these 
unexplored regions. The collections are incor- 
porated in those of Herbarium Bogoriense. Two 
parts of Reinwardtia appeared this year with 
articles on Cryptogams and Phanerogams. Mr. 
Dilmy and Dr. Kostermans attended the UNESCO 
Symposium in Ceylon. 

In 1957, Mr. Forman made a trip to North 
Celebes. Dr. Kostermans collected in East Java, 
especially bamboos. Mr. Dilmy made a trip to 
the U.S.A. and Europe, sponsored by the Ford 
Foundation and UNESCO. Two volumes of 
Reinwardtia appeared. In 1956 more than 
10,000 specimens were sent out in exchange, 
and about 1,000 specimens were received. In 
1957, exchange and loan of material was carried 
on as usual and is still increasing. Dr. Koster- 
mans worked up the Lauraceae of Madagascar, 
which appeared in a series of articles in Bogor 
and Brussels. A monograph on Durio by 
Dr. Kostermans, assisted by a student of the 
Biological Academy, Mr. Sugeng, has been com- 

The botanical activities of the Bureau of 
Horticulture at Pasar Minggu, near Djakarta, 
include the following: 

(1) Distribution of fruit trees. An attempt 
is being made to establish the correlation between 
the distribution of fruit trees with ecological 
factors, viz. rainfall, soil-type, hours of sunshine, 
water-table, temperature. As an example of 
these investigations, it has been found in various 
grape-growing centres that it is very probable 
that a minimum of 70% of sunlight is required 
for success. 

(2) Cabbage breeding. A scheme on the 
breeding of high quality cabbage varieties capable 
of seeding in the tropics is being worked out. 
Crosses between high quality non-seeding varie- 
ties and lower quality seeding varieties have 
been carried out. A vernalisation project on 
non-seeding high quality varieties is being 

(3) Virus free potato seed stock. In connec- 
tion with the problem of keeping a virus-free 
seed stock collection of potatoes, insect free 
chambers are being built in a glasshouse. Various 
plant-indicators are already multiplied for the 



purpose. Import of wild and cultivated species 
of South America is being planned. 

(4) Citrus decline. An investigation into the 
cause of citrus decline is going on. Indications 
are very strong that the decline is not due to 
soil, but to other sources. Nematodes have 
been isolated from roots of plants. Budding 
trials on lime ("djeruk nipis") with scion material 
from declined citrus trees shows stem-pitting and 
vein clearing in the leaves. 

The division of Phytosystematics, -geography, 
and -sociology of the Faculty of Agriculture, 
University of Indonesia, Bogor, put it as one of 
the objectives of botanical investigations by the 
senior students of the Faculty, the study of the 
main features of phytogeography and -ecology 
in Indonesia. Several reconnaissance expeditions 
have been undertaken in 1953 to the Lawu Moun- 
tain (3,300 meter above sealevel) in East-Central 
Java, and Sumatra, in 1954 to Central and East 
Java, Banten (West Java), and the islands in the 
Bay of Djakarta. We have got the opinion that 
in all probability the site factors climate and soil 
series, together with the geographical location, 
have an important influence on the structure and 
floristic composition of our tropical plant com- 
munities. After the departure of the Head of the 
Division, Prof. Dr. S. Bloembergen, at medio 1 955, 
in a very limited scale ecological surveys in trop- 
ical rain forests have been carried out in East 
Kalimentan and the Mount of Lawu. Further 
botanical investigations are planned. Recently, 
Mr. G.A. de Weille, staff member of the General 
Research Station at Medan (North Sumatra), 
has taken his doctor's degree in agriculture 
at the Agricultural Faculty of Bogor, on a thesis 
entitled "Blister blight control in its connection 
with climatic and weather conditions". 

Since 1953 the Central Research Stations 

Association (C.P.V.) at Bogor has published the 

following articles of botanical interest in English: 

(1) Schweizer, J., The physiology of latex as 

basis for tapping systems (1953). 

(2) Wiersum, L.K., Results of some preliminary 

experiments on stimulation of latex 
yields (1953). 

(3) Vollema, J.S. and Lasschuit, J.A., The 

mildew resistant clone LCB 870 (1953). 

(4) Vink, A.P.A. and Alphen de Veer, E.J. van, 

Mechanical and chemical lalang control 

(5) Zeehuizen, J.J. and Arentzen, A.G.J., 

Divergencies between strain and mo- 
dulus determinations (1953). 

(6) Zeehuizen, J.J. and Schoon, Th. G. F., 

Preoculation of the yellow fraction by 
alkaline salts of weak acids with preser- 
vative properties (1953). 

(7) Zeehuizen, J.J. and Schoon, Th. G. F., 

Oxalic acid as a coagulant in the pro- 
cessing of rubber (1953). 

(8) Zeehuizen, J.J. and Schoon, Th. G.F., The 

influence of glyoxal on the vulcaniza- 
tion characteristics of natural rubber 

(9) Wiersum, L.K., Observations on the rooting 

of 1/cvea cuttings (1955). 

(10) Wiersum, L.K., Physiological investigations 

into the process of yield stimulation of 
latex (1955). 

(11) Paardekooper, E.C., Results of testing of 

Hevca seedlings obtained by hand pol- 
lination during 1927-1944 (1956). 

(12) Paardekooper, E.G., The occurrence, pro- 

perties and possible application of 
juvenile type buddings (1956). 

(13) Paardekooper, E.C., Further data on early 

selection in Hevea (1956). 

(14) Knaap, W.P. van der, Results of tea clone 

experiments at Pasir Sarongge (1955). 

(15) Knaap, W.P. van der, Notes on disease 

incidence of Exobasidium vexans Mas- 
see on tea (1955). 




Botany Division, C.S.I.R., Wellington, New Zealand. 


First must be mentioned the death of the 
veteran botanist Dr. W.R.B. Oliver on 16 May 
1957. Dr. Oliver published a revision of the 
genus Aciphylla in October, 1956, (Trans. Roy. 
Soc. N.Z. 84, pp. 1-18), and was active in field 
work until the end of that year, making a collect- 
ing trip to Norfolk Island in November. He 
was able to carry on a limited amount of herba- 
rium work until within a week or two of his 
death. Another loss to New Zealand botany 
was the death, in July, 1957, of Mr, G.O.K. 
Sainsbury who, having completed studies on 
New Zealand and Tasmania mosses, was working 
actively on collections from some Pacific Islands 
and from Malaya. The herbaria of both these 
collectors are at Dominion Museum. 

Distinguished botanical visitors to New Zea- 
land during the years under review included 
Dr. F.W. Went, Dr. H. Godwin, Professor R. D'O. 
Good, Dr. W.M. Hicsey, and Dr. J.W. Gregor, 
all for rather short periods. There was a good 
representation of Australian botanists for the 
A.N.Z.A.A.S. meeting in Dunedin in January, 
1957, and the International Grasslands Congress 
in November, 1956, and the Empire Forestry 
Conference now meeting here have also brought 
numbers of plant scientists to our country. 

Dr. H.H. Allan was one of the twelve biolo- 
gists to receive the honorary degree of Doctor of 
Philosophy at the Linnaean Jubilee celebrations in 
Uppsala in May, 1957. 

With the appointment of Dr. W.R. Philipson 
to the recently established Chair of Botany at 
Canterbury University College, we have, for the 
first time, a full Professor of Botany at each of the 
four main University institutions. 

Dr. G.H. Cunningham, F.R.S., retired from 
the directorship of the Plant Diseases Division 
of D.S.I.R. at the end of September, 1957. 
The name of his successor has not been an- 


Apart from the purely botanical departments 
in the four University Colleges and two of the 
museums, the Botany, Crop Research, Grass- 

lands. Plant Chemistry, and Plant Diseases 
Divisions of D.S.I.R., and the Forest Research 
Institute, increasing amounts of work of direct 
botanical import are coming from several other 
bodies, some of which employ one or more 
professional botanists, e.g., Dominion Physical 
Laboratory, Soil Bureau and N.Z. Oceanogra- 
phic Institute of D.S.I.R., soil conservation 
service under Department of Agriculture, and 
certain local bodies known as Catchment Boards. 

Several rather recently formed societies meeting 
annually discuss botanical problems, e.g., N.Z. 
Genetical Society (established in 1949), N.Z. 
Ecological Society (established in 1951), N.Z. 
Microbiological Society, N.Z. Association of 
Agricultural Scientists, and N.Z. Geological 
Society. The last three formed since 1953. 

In 1953, Botany Division, D.S.I.R., transferred 
to Christchurch, though a small Sub-Station 
is retained temporarily in part of the old quar- 
ters in Wellington. 



In 1956, Chapman published a systematic 
account of the marine Myxophyceae and Chlo- 
rophyceae (Jour. Linn. Soc. 55 No. 360, pp. 333- 
501). Dellow worked on ecology in Auckland; 
Naylor investigated anatomical and life history 
problems in brown seaweeds and produced a 
check list of marine algae for Dunedin. Mason's 
checklist of N.Z. Charophyta appeared in 1956. 
E.A. Flint is currently working on soil algae as 
part of a team dealing also with fungi, bacteria, 
yeasts, amoeboid and ciliate protozoa, and 


In systematics, there has been a steady flow 
of papers from Cunningham and Dingley. G.B. 
Cone has continued her collecting of agarics but 
has not reached publication. A textbook on 
Forest Fungi, prepared by M.E. Lancaster, was 
published for Forest Service in 1955. A study of 
Australasian Cyttariaceae (Rawlings, T.R.S.N.Z. 
84, pp. 19-28) has interests across the Pacific. 



Mycorrhiza and the rhizosphere generally have 
been studied in relation to practical problems. 

Here might be mentioned the work on plant 
viruses at Plant Diseases Division, including 
some very fundamental probing under R.E.F. 


In Dunedin, there is great interest in this group, 
cheifly by Wm. Martin and the chemist J. Murray. 
Unfortunately there has been no N.Z. lichenolo- 
gist amongst the scientists with the N.Z. Antarctic 
party at McMurdo Sound. 


Mrs. Hodgson's work on Hepaticae continues. 
An important landmark was the publication in 
1955, of Sainsbury's Handbook of the New 
Zealand Mosses, with illustrations by Nancy 
M. Adams (Roy. Soc. N.Z. Bull. 5, 490 pp., 

76 pi.). 


Cytological work on ferns is in hand both at 
Auckland and Canterbury University Colleges. 
J.D. Lovis of Leeds spent a year, 1955-1956 
here, continuing his investigation of Asplenium 
trichomanes and comparing southern hemisphere 
forms with those of the north. W.F. Harris' 
Manual of the Spores of New Zealand Pterido- 
phytes appeared in 1955 (D.S.I.R. Bull. 116. 
186 pp., 10 pi. 4 fig.). 


The appearance of Harris' Manual, of Cran- 
well's account of pollen of N.Z. monocots in 
1953, and of Couper's "Upper Mesozoic and 
Cainozoic Spores and Pollen Grains" (N.Z. 
Geological Survey Paleont. Bull. 22, 1953, 

77 pp. 9 pi.) testify to the interest in this field. 
The lack of the dicot pollen atlas promised by 
Cranwell is still acutely felt, but, meanwhile, 
Hards and Moar continue to publish work on 
fresh pollen and that in recent deposits. Atmos- 
pheric pollen studies seem to have been discon- 
tinued meanwhile. Geologists are using plant 
microfossils increasingly for solving chronolo- 
gical problems which still concern us here. 
Questions of the very recent history of the vege- 
tation, e.g., in the last 50,000 years, are also much 
to the fore and C 14 dating services are rather 
generously available. At this very important 
early stage in sketching the outlines of ecological 

change, there seems to have been less than full 
appreciation of Faegri's advice that only a trained 
field botanist will be able to utilise the pollen 
analysis technique fully, and some very poorly 
substantiated statements have been used as evi- 
dence in papers by geologists and physicists. 


Revisions of individual genera continue to 
appear, but Allan's new Flora, Zotov's account 
of the grasses of New Zealand, and Healy's 
manual of the adventive plants in New Zealand 
are all awaited. Preparation is well ahead at 
least with the first of these. 

A noteworthy event has been the flowering 
and fruiting in cultivation of Tecomanthe speciosa, 
plants having been established by cuttings from 
the only known wild plant on Great Island of 
the Three Kings group. 


Experimental and cytological studies are 
helping to clear up taxonomic problems in 
certain groups, e.g., Baylis on Solanum, Brownlie 
on ferns, Connor on Agropyron, Brockie on 
Epilobium, Rattenbury on several genera, and 
Hair on conifers in particular (studies not con- 
fined to N.Z. material) and in general on a 
long-range project of a chromosome atlas of 
New Zealand plants. 


This is mostly on plants of economic impor- 
tance, e.g., crop plants, pasture grasses, and 
timber trees. Bannister's study of a progeny of 
Pinus radiata is giving positive results after only 
six years. Godley is working on dioecy in indig- 
enous plants. 


Lively and varied interest is indicated by these 
titles from A.N.Z.A.A.S. meeting: "Aspects of 
leaf development in Dicotyledons," B.F. Slade; 
"Transformation of vegetative shoot-apex into 
the Primordium of a flower," W.R. Philipson; 
"Effect of various treatments on the Anatomy 
of the stems and roots of apple trees," D.W. 
McKenzie. A paper by R. Licitis-Lindbergs on 
branch abscission and disintegration of the female 



cones of Agathis australis Salisb, (Phytomor- 
phology6, 1956, pp. 151-167) cleared up problems 
that have long awaited attention. 


A most spectacular change in emphasis in 
botanical research here is the increasing interest 
in plant physiology. This does not stem at 
all obviously from leadership in the University, 
but seems to come from practical requirements 
in a country where the national economy is 
based directly on pasture production. This has 
become, for instance, important in the Dominion 
Physical Laboratory, an extract from whose 
current programme reads (D.S.I.R. Annual 
Report, 1957, p. 48) "measurement of mechanical 
properties of plants and the investigation of the 
relationship between these properties and the 
plant's environment; investigation of the moisture 
movement through a soil caused by thermal 
gradients; setting up plant cabinets to provide 
adjustable controlled environments for growing 
plants; development of equipment for the 
measurement of the amount of heat and water 
vapour carried vertically by atmospheric turbu- 
lence; measurement of the transpiration rate of 
plants under different soil moisture conditions 
and in different environments; and, investigation 
of how evaporation over a pasture is influenced 
by pasture length." 

It seems strange that these projects are under- 
taken by a laboratory which has no trained bota- 
nist on its staff. It does, however, work closely 
with Grasslands Division, which is studying 
various other aspects of physiology of pasture 
plants. Basic work on photosynthesis is coming 
from the Plant Chemistry Laboratory. 

Isotopes have been used in various investiga- 
tions and at A.N.Z.A.A.S., Dr. Cone reported 
results using N 15 to support the evidence from 
cultural studies that nitrogen fixation occurs 
quite generally in foliage of pioneer shrubs and 
some weeds and trees. 

Walker and his co-workers have demonstrated 
the importance of added sulphur for optimum 
plant growth in many areas. Jacques' long-term 
studies on roots continue, and there is much 
activity in seed research in relation to pasture 
plants, forest species, and weed control. An 
aspect of plant chemistry that has been given some 
newspaper publicity is a survey of extracts of 
indigenous plants at Auckland University College 
for the Cancer Research Project. 



Descriptive ecological surveys continue to 
hold an important place. The National Forest 
Survey, after ten years of intense activity, has 
reached the stage where a full report on "all 
essential phases of the volumetric survey of 
merchantable-quality indigenous forests" is about 
to be released. The same organisation, within 
the Forest Service, is now proceeding with three 
related projects: (1) Ecological survey to lead to 
regional forest accounts accompanied by mile-to- 
the-inch forest-type maps; (2) Protection forest 
survey covering almost a third of the total land 
area of New Zealand, where the vegetation is 
non-commercial forest, mostly above 2,000 ft. 
altitude, and unoccupied shrublands, grasslands, 
and moorlands that lie above forests; here the 
prime management objective must be regulation 
of the water yield; (3) Exotic-forest survey (F.R.I. 
Annual Report 1957, 24). 

Out of the Forest Survey arose Holloway's 
outstanding paper "Forests and Climates in the 
South Island of New Zealand" (T.R.S.N.Z. 82, 2, 
1954, 329-410). This put forward the hypothesis 
that "the forests, as a whole, are in an unstable 
condition consequent on comparatively recent 
changes in regional climates and that, as a result, 
and active redistribution of species is in progress 
with resultant development of a wide range of 
new, though by no means stable, forest types." 
Evidence is set out to show that the forests of 
the whole of the South Island are comprehensible 
in terms of this hypothesis which is shown not to 
be in conflict with evidence derived through 
studies of the soils or through study of Polynesian 
traditions. This paper is obviously one of the 
highlights of botany in New Zealand in recent 

Regional or local ecological surveys include 
several on tussock grasslands (Barker, A. P.; An 
ecological study of tussock grassland. D.S.I.R. 
Bull. 107, 1953. Tussock Grassland Research 
Committee: The high-altitude snow-tussock 
grassland in the South Island of New Zealand. 
N.ZJ. Set. Tech. 36, 1954. Riney, T.A., and 
Dunbar, G.A.: Criteria for determining Status 
and Trend of High Country grazing lands in the 
South Island of New Zealand. Soil Conserv. 
and Rivers Control Council, 1956. Wardle, P., 
Mark, A.F.: Vegetation and Climate in the 
Dunedin District. T.R.S.N.Z. 84, 1956. Miller, 
F.L. et. al.\ Shotover River Survey. Otago 
Catchment Board Bull. 1. 1956). In this connec- 
tion, Gradwell's studies on frost action in high 



country are relevant. In the North Island, there 
are accounts of forests of Taranaki and West 
Taupo, and of a rather different kind, Druce's 
detailed account of a single catchment of a 
few acres which is to be reserved for long-term 
studies of soil formation and change. 

The "General Survey of the Soils of North 
Island" (1954), the publication of a map showing 
the North Island soil-forming ash showers, and a 
pamphlet on "Soil Pattern in New Zealand" 
will all help plant ecologists. 

Turning to ecology of seashore, Chapman's 
"Mangrove and Salt Marsh flats of the Auckland 
Isthmus" is in the press and intertidal studies 
include Dellow's comprehensive account of 
the shores of the Hauraki Gulf and those of 
G.A. Knox and E.J. Batham in the South Island. 

Field work has not been confined to the main 
island and there have been expeditions to or 
reports on Auckland, Chatham, Three Kings, 
Kermadec, and Pitcairn Islands, and New 
Caledonia. R. Cooper of Auckland Museum was 
working during the winter of 1956 on taro and 
kumara in several parts of the Pacific, and it is 
understood that he has a generous Rockefeller 
grant for similar work further north. 


Members of the Plant Diseases Division 
produced a manual on "Plant Protection in New 
Zealand" (704 pp. 477 illustrations, 1956); and 
in 1954, Chamberlain presented a comprehensive 
account of plant virus diseases in New Zealand. 
At Forest Research Institute, a full-time Forest 
Biology Survey was commenced early in 1956, 
the overall purpose being "to maintain a contin- 
ual and systematic inspection of all forests for 
the presence of newly introduced insects and 
fungi, and to detect any dangerous increases in 
the numbers of existing pathogens." Research 
into the incidence and effects of "noxious animals" 
especially deer and possums, has been stepped up, 
but the teams working on the relations of these 
animals to vegetation are not as strong botanically 

as could be wished. A spectacular drop in the 
number of rabbits (without infection with my- 
xomytosis) has coincided with a series of better- 
than-average seasons, leading to striking vegeta- 
tion changes in some of the more rabbit-ridden 
dry districts. These effects too, have been quite 
inadequately documented owing to the shortage 
of field botanists. 

Manuka blight, a disease of Leptospermum 
caused by a scale insect, has spread partly by 
natural dispersal and partly by deliberate transfer, 
until now, ten years after first being noticed, 
it occurs throughout New Zealand. Affecting 
one of the commonest and most widespread of 
shrub species, it will certainly have direct effects 
on forest regeneration, though it has been 
welcomed by farmers whose poorer pastures have 
always been subject to invasion by manuka. 

A most dangerous addition to the weed flora 
is water hyacinth, which has recently been found 
to seed freely in many waterways and has already 
entailed an expensive control programme. 

On the other side of the picture, plant quaran- 
tine provisions have been much improved with 
the recent installation of modern methyl bromide 
fumigation plants at several main ports. 


At the time of the setting up of the National 
Parks Authority in 1953, there were five National 
Parks, variously controlled. To these have been 
added three more (Mount Cook, 151, 800 acres; 
Nelson Lakes, 139,833 acres; and Urewera 
119,614 acres, and it was enlarged in 1957). 
Moves are afoot to secure reserves in other places, 
especially in Chatham Islands and in the country 
west of Lake Taupo, both regions where active 
agricultural developments have now been made 
possible by use of modern machinery and, in the 
volcanic country, by the use of cobalt to keep 
stock healthy. In both places, the remnants 
of highly characteristic vegetation are dwindling 





Smithsonian Jstitution, Washington, D.C., U.S.A. 

The subcommittee members were selected to 
represent as many countries and branches of 
botany as possible. When completed the commit- 
tee represented continental United States, 6; Ha- 
waii, 2; Japan, 1; Free China, 1; New Zealand, 
2; Australia, 1; and Holland, 1; a total of 14 
members. The majority are taxonomists, but 
phytogcography, ecology, economic botany, 
history of botany, and library science are repre- 

As there was no previous committee on whose 
activities a program could be built, the first activity 
was to circularize the members and a few others 
for their ideas of the possible aims and activities 
of the subcommittee. Following is a summary of 
some of the suggestions submitted. Some members 
pointed out the need for defining more specifically 
boundaries of the area to be included, its present 
limits being a bit indefinite. Severval asked that 
a survey be made of the areas and fields now 
covered by bibliographies, their perspectives and 
adequacy, thus revealing where there are gaps to 
be filled. In order that such information be made 
available to users, it was suggested that a regulary 
appearing bibliographical serial be established. 
Some pointed out the need for a guide for doing 
bibliographical work and the establishment of 
certain standards and procedures. For the benefit 
of workers without access to adequate libraries, 
cooperation between individuals and institutions 
for loans and microfilming, as well as certain 
bibliographical research were suggested. Jt was 
further proposed that editorial assistance and 
guidance be established to aid authors to prepare 
their bibliographic manuscript, so they will be of 
maximum value to researchers and librarians, 
the principal users of bibliographic tools. Close 
cooperation with established bibliographic enter- 
prises, notably the Society for the Bibliography 
of Natural History in England and Biological 
Abstracts, was suggested. An enlarged new 
edition of E. D. Merrill's Bibliography of Pacific 
Botany was proposed. Several of these needs 
would be provided by establishing a bibliographic 
service at some centrally located institution, 
closely linked with similar work at other institu- 
tions, thus broadening the service. Finally, 
careful consideration was urged for the adoption 
of modern new techniques in bibliographic pro- 


cedures so as to broaden their usefulness and to 
conserve limited funds. 

Two progress reports were issued by the sub- 
committee, including summaries of these sugges- 
tions and reports on various known activities 
of the members of the committee and other perti- 
nent endeavors. These were mimeographed and 
distributed by the staff of the Pacific Science 
Association to the members of the subcommittee 
and others, especially to the chairmen of the 
other subcommittees and standing committees, 
and to certain institutions. The present report 
constitutes in effect the third report. 

It may be well here to report on various pro- 
jects in this field. The chairman aided Mrs. Ida 
K. Langman in obtaining a National Science 
Foundation grant which enabled her to spend 
nearly a year searching in Mexican libraries for 
material for her extensive bibliography of Mexican 
botany. She is now extending her work by 
similar searches in American libraries. The 
chairman has rendered some assistance as a 
member of the editorial board for the vegetational 
bibliographies being prepared by Jack McCormick 
at the American Museum of Natural History. 
He reports little recent progress due to other 
obligations. It was found that the Subcommittee 
on Algae is deeply involved in bibliographic work, 
recognizing that this is fundamental to the 
development of any field of study, especially in 
its initial stages. Just how the Subcommittee on 
Bibliography can aid in the bibliographic work 
of the other subcommittees remains to be worked 

For several years Robert Cooper, botanist, and 
Enid Evans, librarian, both at the Auckland 
Museum, have been preparing a bibliography of 
New Zealand botany. Their major problem 
seems to be finding someone with access to 
literature on that area who has the time, ability, 
and resources to run down known-to-exist items 
not available to them. Thus the establishing of 
the bibliographic service proposed above seems 
urgently needed. 

Two most important bibliographies have been 
issued during the existence of this subcommittee. 
First is the bibliography in volume five of the 
Flora Malesiana prepared by Mrs. Van Steenis- 
Kruseman. It will be of inestimable value to 



workers over a wide area. Participation in its 
subsequent improvement has been solicited 
by distributing separately bound interleaved 
reprints to key workers. The second important 
contribution is Island Bibliographies by M.H. 
Sachet and F. R. Fosberg. This is in effect a 
cover title for three separate bibliographies, 
entitled: (1) Annotated Bibliography of Micro- 
nesian Botany, (2) Bibliography of Land Ecology 
and Environment of Coral Atolls, and (3) Selected 
Bibliography of the Vegetation of the Tropical 
Pacific Islands. This was duplicated by the 
National Research Council. 

R.T. Hoogland of Canberra, Australia, has 
revealed that, as opportunity permits in connec- 
tion with work on his New Guinea collections, he 
is accumulating material for a botanical bibliogra- 
phy on that great area. Miss Nancy Burbidge, 
also of Canberra, is accumulating bibliographic 
material in connection with preparation of a 
generic list of Australian plants. Edwin H. Bryan 
of the Bishop Museum maintains several valuable 
bibliographic files on various subjects in the 
Pacific and has submitted two papers for the 
bibliography symposium of the Congress. 

The chairman of this subcommittee, in cooper- 
ation with F.R. Fosberg and M.H. Sachet, and 
under grant from UNESCO, has prepared a manu- 
script supplement to Merrill's Botanical Bibliogra- 
phy of the Islands of the Pacific. Dr. Merrill, 
on retiring from his long and highly productive 
work on Pacific botany, turned over his materials 
accumulated for another edition of his well- 
known Pacific bibliographies to E. H. Walker 
and F. R. Fosberg. The UNESCO grant was 
insufficient and other obligations too great to 
permit the preparation of the supplement that 
will eventually be needed, but this limited work 
will go a long way in the eventual preparation of 
that work. It is in effect a token recognition of 
Dr. Merrill's legacy to Messrs. Walker and 

Work on the needed supplement to the 
Merrill and Walker Bibliography of Eastern 
Asiatic Botany may begin in the near future, 
because of as yet informal encouragement by 
the National Science Foundation and the Ameri- 
can Institute of Biological Science, Encouraging 
promises of full cooperation by Japanese and 
Chinese botanists were received by the chairman 
on his recent visit to those countries. The pro- 
mises are bright that this supplement will better 
meet the needs of the Oriental botanists than did 
the original work. 

Time prevents reporting here on various 
projects that may come within a broader defini- 
tion of bibliographic work. The term is not 
confined to the preparation of bibliographies, 
but embraces all work with books. It would 
scarcely be justified to attempt to enumerate here 
the botanical works that have been or arc being 
prepared in this area or the library developments 
that are pertinent to bibliographic work in its 
widest sense, but such would indeed be very illu- 
minating and profitable. 

In June, 1956, the chairman of this subcom- 
mittee was asked by the Chairman of the Organiz- 
ing Committee on Botany of the Ninth Pacific 
Science Congress to convene a symposium on 
Bibliographic Problems in Natural History in 
the Pacific to be held at this Congress. This 
constitutes the culminating activity of this sub- 
committee. The objective of this symposium will 
be to receive the ideas of its participants through 
a limited number of distributed and delivered 
papers and through free discussion at both formal 
and informal gatherings. The results will then be 
summarized. It is hoped that from the various 
reports of the Subcommittee on Bibliography and 
the coming symposium a more concrete program 
for the development of this work in the Pacific 
may be formulated and implemented. 


F.E. EGLER: I would like to draw your attention to a 
bibliography project now centered at the American 
Museum of Natural History (New York 29). This project 
is essentially the responsibility of Dr. Jack McCormick of 
that institution. It is, or was until recently, entitled 
Regional Vegetation Literature. It is not an all-botany 
bibliography, but one concerned specifically with vegeta- 
tion (plant community) publications with special emphasis 
on identifying the localities at which studies have been 
made. The area involved at present is North America and 
South America, and this is part of our specific area. I 

understand that they are seeking collaborators for different 
countries. Publication of the first of a new series of 
publications is already far advanced. 

F.R. FOSBERG: The supplements to Merrill's Bibliog- 
raphy of the Pacific Islands and to Island Bibliographies 
were aided by a small financial grant by the UNESCO Humid 
Tropics Committee. No provision was made for their 
publication, but there is some indication that funds may be 
available for their publication. 1 would suggest that this 
session could appropriately adopt a recommendation that 



these supplements be published so that they would be of 
help to others. 

M.S. DOTY: The phycology sub-committee has recog- 
nized "bibliography' 1 as perhaps the foremost of the 
barriers to phycological research which might be lowered 
by efforts of the sub-committee. Thus, after consultation 
with Dr. Walker, compilation of bibliographies was begun 
on a form adoptable to his extension of E.D. Merrill's 
Bibliography of Pacific Botany. This material should be 

considered for inclusion in the present "First supplement 
... M to that publication. 

C.G.G.J. VAN STEENIS: Naturally I want to support 
publication of bibliographies. However, it would be very 
desirable to have them in printed form, instead of less well 
surveyable and more bulky mimeograph. Has Dr. 
Fosberg tried to have them published in the Bernice Bishop 
Museum Bulletins or Philippine Journal of Science which 
have a wide circulation in the Pacific area? 






University of Hawaii ', Honolulu, Hawaii. 

Two mimeographed circulars were prepared 
and sent to members of the subcommittee. These 
evoked considerable interest and demand. We 
suggest the practice be continued, but duplication 
and distribution should be done by the Pacific 
Science Association Secretariat in Honolulu. 

Participation in bibliographic matters was 
begun. This was after conferences and corre- 
spondence with the Bibliography Subcommittee 
Chairman and with a local group assembled in 
Honolulu to make a report to that subcommittee. 
In conformation with the style used by E.H. 
Walker of that subcommittee, bibliographic 
materials have begun to be assembled in Honolulu 
regarding algae in the Pacific. Jt is planned that 
this information will be made available to anyone 
at cost or, eventually, be duplicated. We do not 
anticipate duplicating the various efforts to pub- 

lish bibliographic materials but, rather, assist in 
their extension and help make them available 
to prospective users. 

The major phycological problems have been 
sought by correspondence with the subcommittee 
members and enumerated in the mimeographed 
information bulletins distributed to the sub- 
committee. These suggestions were to promote 
floristic studies, develop bibliographic sources, 
facilitate exchange of ideas and materials, and 
encourage functional studies of the algae. 

It is further suggested the Standing Com- 
mittee on Botany of the Pacific pass a resolution 
strongly favoring adherence to the International 
Botanical Code of Nomenclature and in particular 
to those pre-eminent principles the type method 
and priority. 






Pasay City, Philippines. 


The complex problem of common names of 
plants, their variability, recording, and stability 
in some respects, was brought before a session of 
the Eighth Pacific Science Congress. Although 
standardization was undertaken successfully in 
some regions, it was pointed out that standardiza- 
tion of common names of plants in the Pacific 
Basin is not feasible, because too many language 
groups are encountered in this part of the world. 
It was consequently agreed that a subcommittee 
on Common Names of Economic Plants be estab- 
lished, which would work towards the compila- 
tion of a list of common names of plants in 
active use in the Pacific Basin together with their 
botanical equivalent, to be listed in form of a 

As such a vast task can not be accomplished 
uno acttt, the project was divided into the follow- 
ing groups: foodplants, forest trees, weeds and 
forage plants, and ornamentals. For the Ninth 

Pacific Science Congress, only foodplants were 
worked on. 

A list of basic foodplants, based on Philippine 
material, was prepared and sent to botanists 
interested in vernacular names and familiar 
with certain regions in the Pacific. Only generally 
used foodplants were included, not famine 
foodplants or those of local occurrence. Varie- 
ties and hybrids were generally not listed, also 
not all names of products, as those of the coco- 
nut, for instance, could fill a book. 

In the beginning only tropical plants were 
included, but later on it was apparent that even 
in the tropical Philippines many temperate zone 
foodplants are being cultivated and that a separa- 
tion was impractical. In the first attempt, China, 
Japan, and other countries not directly bordering 
the Pacific, such as Thailand, were not included; 
but as the compilation progressed, it became evi- 
dent that such additions might prove very useful, 
as few references are available in Roman script 
in those languages. 


Miss Marie Neal 

Bishop Museum, Honolulu 17, Hawaii. 

Mr. B.E.V. Parham 

Department of Agriculture, Suva, Fiji. 

Mr. R.E. Dwyer 

Director of Agriculture, Papua, Port Moresby. 

Dr. F. Raymond Fosberg 

c/o National Research Council, Washington, U.S.A. 

Dr. Shiu-ying Hu 

Arnold Arboretum, Jamaica Plain, Mass., U.S.A. (since 1957). 

Mr. H. Ando 

Hiroshima University, Sendamachi, Japan (since 1957). 

Mr. Faustino Miranda 

Institute de Bilogia, Casa del Lago, Chapultepec. 

Miss N. Burbidgc 

Canberra, Division of Plant Industry, Australia. 








North Australia. 




Mr. H. Keith 

has resigned on account of transfer. 

(North) Borneo. 

Mr. Jacques Barrau 

Laboratoire d' Agronomic Tropicale, Marseille, France. 


Ida Langmann 
Academy of Science, 
Philadelphia, U.S.A. 

West Coast of North America. 

Dr. Mona Lisa Steiner Philippines. 

2833 Park Avenue, Pasay City, Philippines, chairman. 


As many references as possible were scanned in 
Manila for vernacular names of focdplants in 
the Pacific Basin and combined with published 
and unpublished lists of vernacular names sent by 
members of the committee and others as listed in 
the bibliography. Sincere gratitude is herewith 
expressed here for all the valuable cooperation. 

The Pacific Basin was then divided into various 
regions, such as Micronesia, Polynesia, Malane- 
sia, and the locality or region in which a common 
name is being used is then indicated. Sometimes 
numerous languages are encountered within one 
region, yet frequently one name is of more 
general usage, while others are of local occur- 
rence only. In the Philippines, for instance, one 
language of about 80 has been selected as the 
National Language, namely Tagalog, and this 
language can be understood all over the country. 
The various common names of a region have 
been compiled, but the more generally used name 
has been underlined. In addition to the various 
Malayan, Polynesian languages, etc., the common 
names of foodplants in Chinese, Dutch, English, 
French, German, Japanese, and Spanish were 

Although the chairman is well aware that such 
a compilation can never be complete, an attempt 
has been made to compile all the gathered infor- 
mation in the form of book. With the valuable 
assistance of the National Research Council in 
the Philippines and the aid of Dr. Patrocino 
Valenzuela 200 copies of A PRELIMINARY 
were mimeographed in February, 1957, and 
sent to all members of the committee and 
institutions that might aid in this undertaking. 
Members of the committee have then been asked 

to check the vernacular names given, fill in gaps, 
also to underline the most generally used name 
in a certain region. The preliminary compila- 
tion thus checked and filled in was to be returned 
to the chairman before the Ninth Pacific Science 

Since the compilation was sent out, much new 
material was gathered by the chairman who went 
on six months leave in Europe. Particularly, 
the common names in the major languages were 
filled in. Among those who returned a checked 
copy was Mr. H. Ando, Hiroshima University 
who added more Japanese names. Dr. Shiu-Ying 
Hu, Arnold Arboretum, is preparing Chinese 
equivalents. Other new additions, too, are waiting 
to be included. After the various additions and 
revisions are made and incorporated, it is hoped 
that the first volume of the compilation will 
come out in print. 


1. The arrangement of compiling plants alpha- 
betically within families was more practical for 
the compiler, but should not be followed in the 
final work. The scientific names are to be 
arranged alphabetically. 

2. No personal acknowledgements have been 
made in the preliminary report, but all used 
references, printed and unpublished, were included 
in the bibliography. The final dictionary, however, 
should have special acknowledgements. 

3. So far no cross index has been made, but as 
soon as all the new information has been 
gathered, a cross index should be prepared. 

4. A taxonomist of standing should check the 
validity of the used scientific names and their 



5. Other workers should be appointed to work printing of the revised and enlarged compilation 
on forest trees and medicinal plants while the of common names of foodplants. It should also 
work on foodplants is to be completed. be distributed through the Pacific Science Asso- 

6. A fund should be made available for the ciation. 






Institute for Regional Exploration, University Station, Ann Arbor, Michigan, U.S.A. 

The Subcommittee on Ethnobotany of the 
Standing Committee on Pacific Botany was organ- 
ized in 1954-55 and is composed of twenty-five 

The Chairman has circulated a request for 
information about significant ethnobotanical 
studies currently underway in various parts of 
the Pacific area. Replies have been slow, but 
those that have been received form the basis of 
the following report. 


Dr. Toichi Mabuchi of Tokyo reports that the 
Japanese Society of Ethnology plans to begin 
a field study soon on rice agriculture in Southeast 
Asia, notably in Thailand and Cambodia, which 
will require three years. He is continuing his own 
studies of aboriginal tribes of central Formosa 
and has investigated the correlations between 
social organization and agriculture. Mr. Kokichi 
Segawa, who is now in Brazil for the Japanese 
Government, has been collaborating with Gordon 
Bowles in the compilation of An Illustrated 
Ethnography of the Formosan Aborigines, Vol. I: 
The Yami by Segawa and Kano, and this work 
includes ethnobotanical illustrations and des- 

Thor Heyerdahl has continued his field work in 
the Pacific searching for new evidence for diffu- 
sion. Dr. George F. Carter suggests that the 
discussions at Bangkok "might well include some 
review of this ... with a call for further re-exami- 
nation of the botanical evidence for diffusion." 

Carter urges that botanists and anthropologists 
collaborate to "get that ethnobotanical dic- 
tionary moving, and to include in it meanings or 
names, whenever possible, and to put in the plant 
usages, and most especially ritual usages." 

Dr. Jonathan L. Hartwell, Laboratory of 
Chemical Pharmacology at the U.S. National 
Institutes of Health, is collaborating with various 
colleagues in an extensive survey of the literature, 
both from America and abroad, and folklore, 
pertaining to medicinal plant uses. The purpose 
is primarily pharmacological to open up leads 
for experimental work on the isolation of active 
substances from plants. He is especially interested 
in plants used for the treatment of cancers, 

tumors, warts, and corns, and he requests that 
botanists and anthropologists who are similarly 
interested communicate with him for an exchange 
of information. 

Dr. Harold St. John, who is a subcommittee 
member, will not be at Bangkok but will submit 
papers to two sections, on taxonomy and verna- 
cular names. He is currently in the field for three 
months studying Pandanus in Netherlands New 
Guinea, Australian New Guinea, Papua, and 
Queensland. From this and his other extensive 
work with Pandanus will undoubtedly come future 
ethnobotanical reports. 

Dr. Robert Ornduff writes from Berke^y that 
his current sphere of activity is non-ethnobota- 
nical, that he is doing biosystematic work with 
Senecio species from New Zealand, as well as 
with Californian Composite genera and the genus 
Senecio in the western United States. He is also 
collaborating with anthropologists at Reed 
College in a project on the Warm Springs Indian 
Reservation in Oregon. 

Jacques Barrau has prepared a paper on "Notes 
on the significance of some vernacular names of 
food plants in the South Pacific Islands" which is 
of considerable interest to ethnobotanists. 

R.D. Hoogland writes from Australia that 
"Botanists in Australia have paid very little 
attention to ethnobotany. Apart from the enor- 
mous amount of work still needed in local taxo- 
nomy and the pressure on economic aspects, this 
can probably partly be accounted for by the small 
aboriginal population and the seemingly little 
use made of the native flora." 

"The latter seems to be the case also in New 
Guinea. In comparison with the islands of Indo- 
nesia, where the native flora is extensively used 
in medicine, very little use is made by the New 
Guinea natives of their vast resources in this 
field. During my collecting trips in New Guinea, 
I have tried to collect material of native drugs 
for chemical testing and found it very difficult to 
get any at all. Others have noticed the same, 
e.g., D'Alberts (1880); Professor Baas Becking 
tells me that he has the impression that this 
applies to the whole of Melanesia." 

Anthropologists have been working in various 
areas in New Guinea and have assembled various 



ethnobotanical data, none of which, however, has 
been published in recent years. These data apply 
in particular to food-plants, both cultivated and 
collected in the forest. Others have been working 
in Sumatra and Borneo (from Australia ed.)." 

Robert Suggs is currently doing archaeological 
work in the Marquesas. Previously he indicated 
to Professor Bartlett that he would try to collect 
as much ethnobotanical information as he could 
and would seek native names for the plant spe- 
cimens he hoped to bring back. 

George Carter adds a further note to the effect 
that Heyerdahl's Galapagos expedition "demon- 
strated that the Peruvians had sailed to those 
islands and used them as fishing stations for 
hundreds of years. Since this was presumably 
done with balsas, it strengthens his case that the 
Peruvians were fully capable of sailing into the 
the Pacific and returning... (linguistic similarities 
in Polynesia and South America) the total culture 
picture suggests extremely long continued con- 
tacts, with some of the later ones coming from 

Jonathan Sauer writes from the University of 
Wisconsin that he is continuing his work with 
grain amaranths, and with Sasuke Nakao has 
published further data and interpretations of 
Asiatic grain amaranths, crops domesticated by 
the American Indians and introduced to Asia by 
unknown means (Land and Crops of the Nepal 
Himalaya, ed. H, Kihara, pp. 141-146. Kyoto, 
1956). ' 

Dr. Sauer is also continuing to grow specimens 
of pot-herb amaranths collected in seed from 
various parts of Southeastern Asia, preparatory 
to systematic study, and he reports that he is 
currently engaged in a monographic study of all 
species of Canavalia (sord-bean and jack-bean) 
from all parts of the world, with special emphasis 
on the aboriginal cultivated varieties. He invites 
workers to send him specimens of Amaranthus 
or Canavalia and promises to identify them and 
give a report to the collectors. 

Dr. Tom Harrisson indicates that he has col- 
lected considerable information on the eth- 
no -ecological aspects of primitive agriculture in 
Borneo and that he is willing to exchange infor- 
mation. He is reporting on ethnobiological lore 
in Borneo at the Congress. 

Harold C. Conklin is currently in the Philip- 
pines to study tropical forest agriculture and will 
present a number of papers at the Bangkok 

Professor Bartlett, now an emeritus professor 
at Michigan, is principally engaged in compiling 


a gigantic annotated bibliography on the subject 
Fire in Relation to Primitive Agriculture and 
Grazing in the Tropics. Volume 1 has already 
been distributed and is about out of print; volume 
2 is out and ready for distribution (A-G, Tropics 
in general; H-J, South Asia and Oceania a total 
of 873 pages); and he has enough material already 
gathered for a third volume of equal proportions. 
He is also actively advising ethnobotanists, 
archaeologists, and botanists who are in the field 
or about to take to the field as to what problems 
still need to be attacked. He writes that "sweet 
potatoes introduced into the Orient by the Spanish 
should be compared in every way, cytologically, 
chromatographically, and otherwise, with those 
that spread into Polynesia from New Zealand." 
He is especially interested in the possibilities of 
tracing sweet potato migrations by means of 
chromatography. He also wants someone to 
study native gardening carefully all along the 
Southeast Asia coast. 

As an appendix, a brief report is attached 
covering studies on Manihot esculenta by Dr. 
David Rogers who is continuing this work at the 
New York Botanical Garden, in addition to 
editing Economic Botany. 


In Hokkaido, Japan, Dr. Robert Austerlitz 
has been working for several seasons with Gilyak 
and Ainu informants. His project is primarily 
a linguistic study, but he has developed an active 
interest in ethnobotany and has compiled a large 
annotated list of plants used for foods, medicines, 
poisons, and handicrafts. His specimens and 
field identifications have been checked by Dr. M. 
Tatewaki of the Botanical Institute at the Hok- 
kaido University. 

T.P. Bank II is continuing studies of Aleut and 
Pacific Eskimo plant lore. Recently he spent 
a year in Hokkaido to compare this material with 
what is known about Ainu plant uses. He is 
especially interested in Eskimo- Aleut migrations, 
Aleut medical knowledge, and the question of 
cultural diffusion across the North Pacific. 

Bank is commencing an ethnobotanical and 
toxicological study of aconite poison whaling in 
the North Pacific, in collaboration with workers 
at Hokkaido University. Part of the study will 
include an evaluation of the variation of toxicity 
in various races of Aconitum and the significance 
of this to anthropological questions pertaining to 
the use of aconite poison by primitives in the 
North Pacific region. 



Archaeological studies by Bank and others at 
Michigan have also utilized ethnobotanical 
methods for determining chronology of cul- 
tural events in the Aleutians. Radiocarbon 
dating of archaeological deposits and pollen 
analysis have indicated that the Aleuts arrived in 
the Aleutians at least 3,500 years ago at a time 
when the climate may have been appreciably 
colder. Preliminary results of pollen studies show 
that the beginning of amelioration of climate 
toward the onset of the thermal maximum in the 
Aleutians may have occurred between 4,900 

and 4,000 years ago. 

Others are doing botanical and ecological work 
in the Aleutian area, including Robert Kindschy, 
Jr. at the University of Idaho, who collected 
ethnobotanical data among the Nikolskian Aleuts 
in 1957. 

No reports have been received from ethno- 
botanists at the University of Washington, but 
presumably studies are being continued among 
the Indians of that region and in southeastern 





New York Botanical Garden, New York, U.S.A. 


1. To provide a convenient classification of 
the cultivars of Manihot esculenta. 

2. To demonstrate relationships among the 


1. Assemble cultivars in museum plots for 
ready study of the varieties. 

2. Classify, using morphological character- 
istics, in a framework of taxonomic categories. 


Jamaican and Central American cultivars represent the 
range of material so far studied. In Jamaica, a collection 
of 107 named varieties were brought together from all 
parts of the island at Bodies Experimental Farms. After 
considerable study, forty-five cultivars were recognized. 
In Costa Rica, at the Inter-American Institute of Agri- 
cultural Sciences at Turrialba, Dr. Jorge Leon had collected 
the locally occurring cultivars. To this collection were 
added variants from other Central American countries, 
Cuba, a few from the East Indies, and several from Brazil. 
In this collection about twenty-five cultivars are found to 
be distinct. 

The cultivars from both areas fit into a framework of 
sub-specific categories of two convariants (equivalent to 
subspecies) and several sub-convariants (comparable to 

The classic divisions of Manihot esculenta (sweet vs. 
bitter) have been set aside in this work. No morpholog- 
ical categories are coordinate with such a division. 

Work is in progress to publish a key to the cultivars 
studied, and proposals for continuation of the studies in 
South America are being made. 






The distinctions between ethnobotany on the 
one hand and economic botany on the other are 
not always clear. This has been brought out 
during the present Congress, where papers repre- 
senting both points of view are presented in a 
symposium on the ethnobotany of Thailand and 
contiguous countries. Indeed, ethnobotanical 
reports are distributed throughout various ses- 
sions dealing with a number of diverse disciplines, 
including anthropology, botany, linguistics, 
conservation, and agriculture. This is not a 
deadly sin, but it does make for a lack of coordi- 
nation and a scattering of thoughts and conclu- 
sions, when perhaps the true function of the study 
of ethnobotany is to bring them together. 

Ethnobotany should deal with verbal traditions 
only, except for those traditions which have 
recently found their way into the literature, 
whereas economic botany is more concerned with 
the modern concepts of the science of plants and 
of their uses. The word "primitive" has various 
connotations among different people, but perhaps 
it is still our best key to the content of ethno- 
botanical study. Ethnobotany is the study of 
the interrelations of primitive man and plants, in 
the sense that primitive denotes a lack of any 
written language and therefore the preservation 
of traditions by verbal means only. 

Chemical analysis of useful plants is not usually 
considered a part of ethnobotanical work, but 
the study of plant debris in archaeological depo- 
sits, studies of plant patterns in the vicinity of old 
village sites, pollen analysis, and radiocarbon 
dating are all related to our main objectives in the 
study of ethnobotany. 

With this rather brief definition of the content 
of pur study, the following specific recommen- 
dations are made in the hope that they will stimu- 
late a more thorough consideration of how 
ethnobotanical research can contribute most to 
the solution of problems in the various disciplines 
of which ethnobotany is a part. 

1. A summary of our state of knowledge of 
Southeast Asian ethnobotany is clearly needed. 
It is important that we should not leave ethnobo- 
tany in its present unorganized state of dispersed 
data throughout monographs which are actually 
in content anthropological, botanical, geograph- 
ical, linguistical, or agricultural. For example, 

a great deal of plant lore is included in folklore 
and written literature of Southeast Asia but was 
nevertheless not put there for the purpose of 
transmitting the oral traditions pertaining to 
plants; it includes scattered references to plant 
uses compiled by casual travelers, biologists, 
linguists, and others. Often, this material is not 
the sort that is recovered by Occidental scholars 
but, if found, is more likely to be assembled by 
native scholars. One objective of Thai ethno- 
botanists might reasonably be to round out a 
regional monograph on ethnobotany that would 
stress man's relationship (traditional) to the 
environment through trial and error. 

2. A matter that is frequently neglected in 
making dictionaries is adequate attention to 
primitive gardening and agriculture. Although 
there are dictionaries of primitive tribes which 
have valuable ethnographic content, there are 
many languages that have no records of plant 
names, implements used in gardening, and so 
forth. A comparative study of gardening in 
Eastern Asia has not been made, but such a 
monograph would be extremely useful. 

3. Because of its inter-disciplinary nature, 
ethnobotanical study is frequently pursued by 
persons with a variety of training, with a result 
that many different field methods are employed, 
and the data are not presented in any uniform 
manner. The effective worker should be enough 
of a botanist to make careful and systematic 
collections for identification of the plants and 
precise meaning of the words. Nothing is more 
frequently commented upon by botanists than 
that the materials collected for identification by 
non-botanists are too often hopelessly inadequate. 
Every contributor to ethnobotany, whether he be 
a linguist, anthropologist, or geographer, should 
become enough of a botanist to discriminate 
critically between related plants and to have at 
least as discerning an eye as one who is frequently 
referred to somewhat condescendingly as a 
tribesman. The ethnobotanical worker cannot 
always be a linguist, but he should equip himself 
with at least the basic knowledge of approved 
linguistic field recording techniques and train 
himself to distinguish at least the major sound 
units of the language with which he is working. 
He should of course attempt to fit his botanical 
data into a cultural context, which presupposes 
at least a basic knowledge of the culture he is 



entering and a strong curiosity about the part 
*vhich plants play in this culture. 

4. International cooperation is indispensable. 
\uthorities are not numerous for most plant 
groups; linguistic information for many tribes is 
;carce. The field worker is faced with the dis- 
:ouraging task of either identifying everything 
limself with the help of plant keys, which are 
lard to find, or else seeking out the appropriate 
jpecialists and requesting their cooperation. 
It would be of inestimable service to field workers 
f rosters of specialists could be compiled for 
^ach major region in the Pacific Basin and for 
Southeast Asia. This is a service that could be 
Derformed by the ethnobotanists themselves. 

5. The program in ethnobotany of the Pacific 
Science Congress might best be used as a basis 
or indicating what might be done by scholars in 
5iam and elsewhere, who themselves may not 
lave an adequate means of securing identifications 
)f all materials but who because of their linguistic 
ibilities with vernacular names are in a strategic 
position to cooperate with international agencies. 
\s a step in this direction, a list of suggested 
esearch, compiled by Professor H.H. Bartlett 
U.S.A.), is appended. 

6. One phase of cthnobotanical endeavor that 
las received but little attention in Southeast Asia 
s the securing of chronological correlations with 
)lant materials preserved at or near archaeolog- 
cal sites. Although pollen analysis might not 
)e feasable in some parts of the tropics, because 
>f poor preservation of the pollen, it nevertheless 
ihould be attempted. Dendrochronology may 
ilso prove practical in some areas. Ethnobot- 
inists should be encouraged to work with the 
dentification of plant materials from archaeolog- 
cal sites and seek comparative ethnographic 
:orrelations for their material. They can and 
hould contribute to the rapidly growing collec- 
ions of plant materials for radiocarbon dating 
>f cultural and biological events, and especially 
hey could be of service in insuring that the proper 
>recautions are taken during collection of the 
naterials so that contamination from biological 
ources is avoided. 


A List of Suggested Ethnobotanical Research 
or Southeast Asia Compiled by Professor H.H. 
3artlett, University of Michigan, Ann Arbor, 
Michigan, U.S.A. 

1. Listing of plants mentioned in old Siamese 

vernacular literature, with notes on identity, 
utility, ceremonial importance, etc. 

2. Listing of plants mentioned in old Siamese 
translations of Indie Buddhist literature, etc., 
with parallel listing of Siamese and Tndic (San- 
skrit, Pali, Prakrit) plant names. 

3. Notes applying to specific geographic locali- 
ties on the utilization by the Siamese of local 
plants of all sorts. 

4. Descriptions, for specific localities, of 
ancient traditional Siamese horticulture, village 
gardening, industrial preparation, marketing and 
utilization of plant products, especially indigo 
and other dye plants, palm sugar, starch, glue, 
tannin, etc. 

5. Descriptions, with local names, and utili- 
zation of the cultivated or wild varieties of any of 
the following: bamboo, rattan, upland rice, yam, 
other root crops, coconut, pot herbs ("greens"), 
spices, narcotic plants (for example, betel nut 
palm; betel pepper), fruit trees, etc. 

6. Materials or house-building botanical 
source, geographic origin, mode of transpor- 
tation, etc. 

7. Plants traditionally used in medicine, and 
the lore concerning them. 

8. History of the ancient pharmacopoeia. 

9. Plant poisons and the lore concerning them, 
with reference to vernacular literature. (Control 
of insect pests, rodents, rice birds, predatory 
animals, etc.) 

10. Descriptions of primitive agriculture of any 
localized ethnic or linguistic minority groups, and 
of their village gardening, with notes on specific 
plants, vernacular names, utilization, etc. 

11. Description of sacred trees and groves, and 
botanical identification of the species concerned. 

12. Ceremonial uses of plants. Use of plants in 
religious rituals. 

13. Traditional ornamental gardening in specific 

14. Materials used in basketry. 

1 5. Textile plants ; history, traditional utilization. 

16. Utilization of seaweeds for food or other 

17. The plant lore of weeds and wild plants. 

18. Effect of population impact on remaining 
forest areas in Siam; extent of man-made grass- 
lands; utilization of same for grazing, etc. 

19. History and method of use of lontar palm 
leaves as writing material in Thailand, illustrated 
by lontar manuscripts. 



20. Ancient methods of making indigo (and 
other dyes) with information on the plants used. 

21. The geography, mode of growth, harvesting, 
and utilization of floating rice in deep flood-waters 
of Siam. 

22. The sources and preparation of spices and 
perfume used by local tribes. 

23. The varieties, cultivation and utilization of 
Pandanus in Thailand. 

24. The process of manufacturing sugar from 
the sugar palm (or other source). 

25. The enrichment of abandoned dry rice lands 
with useful woody plants by aboriginal tribes. 

26. The useful plants of the Semang (or other 
people) of Thailand. 

27. Sacred forests in Thailand. 

28. Sacred trees in Thailand. 

29. The use of fungi as food in Thailand. 

30. Food plants of the silkworm in Thailand. 

31. The identification and use offish-poisoning 
plants in Thailand. 

32. Native systems of classifying plants. 

33. Plants used as sources of drinking water. 

34. Plants producing gums, resin, and rubber, 
and methods of utilization of the products. 

35. Primitive methods of making fermented 

36. Plants used as emergency and famine foods. 

37. Village horticulture. 

38. Plants used for thatch. 




Compiled from letter by H.J. Lam of Rijksherbarium, Leiden, Netherlands. 

Since the last Pacific Science Congress in 1953, 
the chairman of the subcommittee has been 
unable to devote a significant amount of time to 
the project of preparation of distribution maps 
which was the task of this subcommittee. 

The Philippine Journal of Science has offered 
to publish the maps and accompanying text, if 
it were carried to a point where this is indicated. 
The main problem outstanding seems to be that 
of securing funds to employ a competent bota- 

nist to do the actual work of compiling the maps. 
Prof. Dr. S. Bloembergen, formerly of Java, is 
known to be available for this task, if funds, to 
the extent of $1,000 per year for half time, or 
$3,500 per year full time, can be made available. 

Office space and facilities in the Rijksherba- 
rium, as well as the close cooperation of Dr. Lam, 
will be made available. 

It is urged that efforts be made to find the 
necessary funds to carry this project forward. 






Gothenberg, Sweden. 

During the Seventh Pacific Congress, Auckland, 
N.Z., 1949, 1 resigned as Secretary to the "Stand- 
ing Committee for the Protection of Nature in 
and around the Pacific" and was succeeded by 
Dr. F.R. Fosberg. During the Eighth Congress, 
Manila, 1953, the Standing Committee for Pacific 
Botany was organized with Dr. Fosberg as Chair- 
man and Mr. Kasin Suvatabundhu as Secretary, 
while I was to take charge of a Subcommittee on 
Nature Protection. Of the former local represen- 
tatives few could be counted on any longer so 
that new members had to be found. 


As was pointed out on former occasions, the 
Imperial Japanese Government has long been 
actively protecting the natural resources of Japan, 
and there has been no reason for the Pacific 
Congresses to make recommendations. Through 
the Japan Nature Protection Society, the general 
public is engaged in the movement, and among 
the directors and trustees are many representa- 
tives of cooperative bodies such as the National 
Parks Association, the Ornithological Society, 
the Forestry Association, Landscape Society, 
Alpine Club, etc. 

(DR. QUISUMBING in letter to 

PROF. VAN STEENIS, Dec. 10, 1954). 

"There are three categories that could be in- 
cluded in the matter of conservation, areas or 
localities, rare species and interesting species. 
The matter... is really a very serious one... . One 
of the greatest difficulties that we are encountering 
in the Philippines is the matter of kaingin (shifting 
agriculture) and also the vast cutting for the lumber 
industry. Many of the areas... cut for timber 
have been declared agricultural lands, to place 
people from congested areas... . Doubtless in 
this project many species are liable to be destroyed 
and perhaps disappear for good. A national 
committee has been appointed by the President of 
the Republic to reserve certain forests... and we 
have many reservations now. The difficulty ... 
is that cutting for kaingin purposes continues and 
it is very difficult to police these areas for lack 
of forest guards and on account of the fact that 

these people in the remote barrios need a place 
where they can expand and cultivate crops even 

Conservation of areas. "In Mindanao we have 
a sizeable area covered by Lansium domesticum. 
The Japanese had killed some of these trees during 
the occupation and some of the natives in that 
particular area went as far as killing trees to 
collect the fruits... . I have indicated to our autho- 
rities that the area should be preserved. In another 
area in the province of Zambales, Vanda Lamella- 
ta var. boxalli is very common on trees. I have 
seen this place before devastation... with the trees 
covered by this orchid in flower. As most of 
these areas are private lands we find difficulties in 
having them preserved. Even if the government 
would expropriate the land, it is difficult to con- 
trol the vandals; the orchids are collected in bud 
and brought to Manila by peddlers. There is also 
a project now of conserving strips of forests in the 
central portion of the island of Leyte. 

I am going to submit a list of plant species 
included in the law for conservation, but as my 
files have been destroyed I shall look into the 
original law to get the names. One is Lilium 
philippinense, which was very common in the 
grassy area in the pine belt of mountain Province. 
In early days it was indeed a sight to see this 
lily when in bloom. The Igorots have been 
gathering plants and brought to Baguio for sale. 
...Another species in the list for conservation is 
Phoenix hanceana var. philippinensis which is 
found only on the island of Pabtan. The main 
species is reported from Southeastern China, 
Thailand, and Formosa." 


(according to letters to PROF. VAN STEENIS). 

Malaya. J. Wyatt-Smith, Forest Botanist, 
Kepong. Oct. 11, 1954. 

"The position as I see it is that it is impossible 
with the present botanical knowledge of this 
region to consider reservation for the preservation 
of isolated species or endemics except in very 
special circumstances. Future collections, explora- 
tion and taxonomic studies may well reveal the 
occurrence in abundance of a hitherto "rare" 
species and endemics may no longer be endemics 



through being found in other countries or by 
being reduced in taxonomic studies. Further, 
if I am not committing botanical heresy, is 
it so vital to preserve isolated endemic species 
in a genus unless there is something very special 
about them which is of interest phylogenetically ? 
Endemic genera and single representatives in a 
country or outliers of a genus by all means." 

"I am wholeheartedly in support of preserving 
large areas of all forest types throughout the 
country, and thereby preserving complete asso- 
ciations together with the rest of the plants and 
animals, etc., making up the synusia. With this 
in mind several Jungle Reserves have already 
been constituted in Malaya and many more are 
under discussion (see Malayan Forester, Vol. 
XIII, 1950, pp. 92-94, and Malayan Annual 
Reports), and steps are also being taken to 
preserve certain areas such as Jugra Hill which is 
the only place in the Federation where Shorea 
gratissima is found." 

"As regards preservation of 'existing isolated 
rare' species..., the Malayan Nature Society is a 
fairly influential body and preservation and 
conservation has already come under discussion. 
I am Secretary of the Selangor branch and closely 
connected with the main committee... so I am in 
some sort of position to push anything you con- 
sider desirable in this connection." 

Singapore Island. J.W. Purseglove, Director, 
Botanic Gardens, Singapore. 

"I am in complete agreement that everything 
possible should be done to preserve plant species 
which are in danger of extinction and 1 will do 
all 1 can to help in this matter. The total area of 
Nature Reserves in Singapore is approximately 
8,500 acres controlled by a Nature Reserve Board 
of Management of which I am ex-officio Chair- 
man. The area reserved is a noteworthy achieve- 
ment of a small colony of some 224 sq. miles with 
a population of over 1 million. T doubt if we 
can hope for any further reservation on this 

Further information on Singapore Island is 
given in a circular "Nature conservation on 
Singapore Island and in particular the threat to 
the Pandan Nature Reserve", signed by H. 
Burkill, Chairman, Nature Reserve Board, Botan- 
ic Gardens, Singapore, 1957. The circular gives 
the history of the forest reserves, which are 
characterized as follows. 

"Each of Singapore's Reserves, besides the 
general purposes of recreation and education, 
was created for a particular biological reason. 


Rare plants exist in all of them. Bukit Timah 
is the classical collecting area for a very large 
proportion of Malaya's flora. It is the type 
locality for probably more species of plants than 
any other area of comparable size in the world. 
It is the only remaining piece of primary lowland 
forest on Singapore Island. Kranje typifies the 
transition from muddy fore-shore to a dry land 
vegetation, whereas Pandan is typical tidal 
mangrove. These two form a series in the land- 
building character of mangrove vegetation. With 
the total exploitation of other mangrove areas in 
South Johore, Pandan is the last remaining piece 
of typical mangrove. It is moreover the most 
southerly of continental Asia. Labrador conserves 
a primitive species of fern, Dipteris conjugata, 
which is found elsewhere in Malaya only above 
3,000 ft. altitude. Pandan is the type locality 
for many marine animals and all the Reserves 
and the Water Catchment Area are sanctuaries 
for wild animals." 

Of great general interest is Mr. Burkill's answer 
to the proposals to utilize Pandan: 

"Proposals have been made to clear the whole 
of the Pandan Reserve for the establishment 
of prawn ponds. This is the second attempt to 
remove control of the area from the Nature 
Reserves Board for commercial exploitation, 
the previous one in 1956 being successfully 
resisted. It is not the purpose of these notes to 
set out the arguments for these proposals. Suffice 
it to say that the Nature Reserves Board is un- 
willing to cede the area since it is a legally con- 
stituted body with a duty to perform." 

The following general remarks regarding 
Malaya may also be quoted from Mr. Purse- 
glove's letter to Dr. van Steenis (see above). 

"With the present state of collecting within 
the Malaysian region, are we justified in assum- 
ing that a plant is in danger of extinction because 
it has only been found in one small area? Until 
much more collecting has been done and the 
flora and its distribution known more fully than 
at present, the possibility of rare and interesting 
species or species at present only recorded from 
one small locality turning up in an area at present 
little collected would seem to be considerable. 
1 think that the most we can do at the moment 
is to list those species which reputable botanists 
consider in danger of extinction and do what 
we can to preserve them, the method of preserva- 
tion being dependent upon local conditions." 

"Unless a sufficiently large area is preserved 
ecological conditions will be altered which might 



result in the loss of species. In secondary com- 
munities reservation may result in a change in 
vegetation with resulting loss, especially when a 
plant may depend for its continuance on biotic 
factors. It would be difficult to look after a 
large number of small areas and see that they are 
adequately patrolled, even if money and staff 
are available. Making a small reserve without 
adequate patrol and advertising the presence of 
the species might lead to its more rapid extinction. 
The possibility of preserving the plant in cul- 
tivation might be considered." 


In addition to the late Dr. K.W. Dammerman's 
beautifully illustrated work, "Preservation of 
Wild Life and Nature Reserves in the Nether- 
lands-Indies," published for the Fourth Pacific 
Science Congress in 1929, comprehensive reports 
appear in the Proceedings of the Fifth Congress, 
Vol I, 1933, but after that no further report was 
received until 1949, when a few remarks by Dr. A. 
Hoogerwerf were included in the Proceedings, 
Vol. IV, 1953, referring to the work done by the 
Netherlands Indies Association for Nature Protec- 
tion which issued annual reports. 

It would have been of particular interest to 
obtain official statements as to what happened 
during the Japanese occupation and the following 
insurrection which ultimately led to the establish- 
ment of the Republic of Indonesia. It is, however, 
good to learn from a letter to Dr. van Steenis, 
written Oct. 25, 1954 by the Director of Kebun 
Raya Indonesia (formerly Buitenzorg Botanical 
Garden), Prof. I. Kusnoto, that the efforts made 
by his predecessors to convince the government 
of the necessity of plant protection by law will 
be renewed, and that everything will be done in 
order to intensify plant conservation. A restric- 
tion on the export of rare species, among them 
orchids, is planned, as well as the protection of 
the localities for rare species, by setting aside such 
areas as reserves. For detailed information the 
departmental Head of the Herb. Bogoriense and 
the Chief of the Botany Division of the Forest 
Research Station should be approached. 

Sumatra. Dr. W. Meijer recently published a 
paper on "Natuurbescherming in Midden-Su- 
matra" (Natuur en Landschap 101 , 1956), contain- 
ing an interesting reconnaissance of protected 
natural monuments such as Harankloof, a low- 
land virgin forest at Pantji conserved since 1930 
and locality for Rafflesia Arnoldi, Aneikloof, 
type locality for Amorphophallus titanum, etc. 

The forest reserve on Mount Sago, 2,000 meters, 
is now being investigated scientifically. An area 
of 10 hectares, 1,000 m. above sea level, included 
150 different kinds of trees, almost equalling 
Tjibodas on Java in richness. Nature conser- 
vation in Sumatra is an enormous task, but 
wholeheartedly supported by the Governor and 
the Director of the Forest Service. The immensely 
interesting extinct volcano Mt. Singgalang has 
recently been proposed as a forest reserve. 


(The late G.H.S. WOOD, Forest Botanist, 
in letter to DR. VAN STHENIS, Oct. 5, 1954). 

"Regarding the preservation of species and 
ecological types I am following Malayan practice. 
This is primarily the making within forest reserves 
of virgin jungle reserves of 50 to 100 acres which 
will not be interfered with in any way. Con- 
stituted forest reserves at present occupy only 
3% of the total land area of North Borneo, but 
it is anticipated that large areas now being worked 
as concessions will become forest reserves. No 
jungle reserves have so far been made but it is 
expected that creation of such reserves will be 
made obligatory on concession areas and con- 
stituted reserves. The object is to keep samples 
of ecological and physiognomic types rather 
than particular species undisturbed." 

"Forest reserves are also being made in forest 
stands which are not necessarily economic (or for- 
seeablyso); hence numerous species are being 
preserved intentionally or not. An example of 
this kind is Tabawan Island in Darvel Bay, which 
I recently visited with Mr. Wyatt-Smith. This 
island supports a peculiar forest type dominated 
by Casuarina sumatrana, Calophyllum spp. and 
an unidentified tree, known locally as bangkan; 
only three scarce Dipterocarpaceae are recorded 
so far." 

"You will appreciate that in general forests 
on the flat land of the east coast of North Borneo 
are undisturbed and offer considerable scope for 
retention as forest reserves containing selected 
virgin jungle reserves. On the west coast, both 
flat land and mountainous slopes up to 3,500 ft. 
have suffered severely from either permanent or 
shifting cultivation. Undoubtedly some species 
have already become extinct and more will follow, 
and little can be done to protect individual species. 
But priority is given to the formation of forest 
reserves on the west coast before the remaining 
virgin stands, particularly at low altitudes, are 
destroyed; hence the great majority of species 



will survive apart from those which are very 

"It is expected that it will be easy to reserve 
areas over 3,500 ft. on mountains on the west 
coast. A Land Utilization Committee is planned 
for North Borneo and I shall propose that the 
whole of Mt. Kinabalu-Mt. Tambuyukon area 
and the Mt. Trus Madi area should be reserved 
as national parks (game and forest reserves) quite 
apart from the formation of other forest reserves 
in the mountains. Proposals have already been 
made for a national park in the Ulu Segama area. 
On the other hand little can be done on the 
flat coastal fringe of the west coast which is now 
relatively densely populated. In this area few 
untouched samples of lowland forest remain on 
well-drained ground, and existing swamp forest 
(either in forest reserves or outside) is threatened 
with conversion to padi land. Generally speak- 
ing we have so little information on the distribu- 
tion of rare or local species that even if it were 
economically feasible to make reserves for them 
one would not know where to make them. In 
the case of tree species it is easier, for there is 
justification in preserving them as part of a partic- 
ular forest type and very often this means that 
the rest of the distinctive associated synusium of 
lianes, under-storey trees, and shrubs is also pre- 
served. In this connection you may be interested 
to know that most limestone hills on the east 
coast (with their economic importance as pro- 
ducers of birds' nests, which are controlled by the 
Forest Department) are being included within 
forest reserves. As we become acquainted with 
other ecological types I earnestly hope that they 
can be similarly treated, but no one can deny 
that large areas at present under forest in North 
Borneo must ultimately be lost to permanent 
agricultural land as the population increases. 
Only careful choice of forest reserves can preserve 
most of the species." 

(August 3, 1955.) "It has occurred to me that 
the project suggested in your letter of March 
8, 1955, might be easily extended by enlisting the 
aid of either FAO or UNESCO, the former 
being particularly active in Malaysia. I know the 
idea is not new but some sort of census or survey 
of existing or potential areas of purely scientific 
value would be of great and permanent value. 
To start with, some idea of the extent of such 
areas might be obtained, followed by detailed 
surveys with estimation of animal and plant 
populations. Being of primarily scientific turn 
of mind, I am particularly interested in such a 
project and I am sure the aid of many scientific 


bodies could be brought to bear on the preserva- 
tion of species which would otherwise be liable 
to extinction." 

A few days later (August 9, 1955), Mr. Wood 
sent a circular letter to seven timber trading com- 
panies of North Borneo. 


Reservatjon of virgin junglej>lots. 

This matter was raised at a meeting with repre- 
sentatives of the main timber firms in Sandakan 
on 6.8.55. A number of Forest Reserves have 
been constituted, which help the preservation of 
much of the native flora and fauna of North 
Borneo, but it is desirable to make as many of 
these as possible in widely scattered areas even 
if they are small. In concessions concessionaires 
are asked to provide details of any areas which 
they do not wish to exploit (e.g. inaccessible 
areas on hills, swampy areas, etc.). These in- 
accessible areas are often the home of rare plants. 
On his recent tour Professor H.G. Champion 
said that such virgin jungle reserves should not 
be less than 50 acres, with a substantial surround; 
further details can be found in Chapter XIV 
of the North Borneo Forest Manual. Naturally 
reserved areas of this sort are just as desirable in 
forest which is commercial and provide a yard- 
stick for regeneration procedure in adjacent 
exploited areas. The reservation of virgin 
forest in North Borneo will allow the preservation 
of most native species, especially in the national 
parks on Mt. Kinabalu and in Ulu Segama, but 
additional widely scattered small areas of virgin 
jungle left in concession are of great value." 

"Please send lists of any areas of this sort which 
you wish to abandon under the terms of your 

(November 26, 1955.) "Further to my letter of 
3.8.55... another point has occurred to me re- 
garding the conservation of rare species and veg- 
etation types. You may be aware that present day 
silvicultural treatment in forest reserves through- 
out the tropical belt of the Commonwealth are 
tending more and more to the elimination of 
weed trees by poisoning. From a scientific point 
of view this is serious, because many of the weed 
trees include rare species which one wishes to 
protect. If forest is logged but not silviculturally 
treated, there seems to be little reason why such 
species should not persist, but systematic poison- 
ing may eliminate them in the long run." 



"I have a feeling that large climbers would be 
more affected than under-storey trees, for not 
only are they broken when large commercial 
trees are felled, but they have to be ruthlessly 
poisoned if they are not to inhibit the young 
commercial saplings of the new crop. If this is 
the case one can foresee a time when forest 
reserves (other than protection and ecological 
reserves) would be largely composed of com- 
mercial trees alone." 

"I may be wrong in my assessment of the recu- 
perative powers of the under-storey trees, but 
quite frequently one meets examples of species 
which are so rare in the restricted localities in 
which they do occur that I hesitate to predict 
their future in the event of intensive silvicultural 

"It strikes me that this problem is much worse 
in Malaysia than in Africa, for here we have so 
many endemic species of limited range." 



c/o The Museum, Kuching, in letter to 
DR. VAN STEENIS, October 13, 1954). 

"1 have consulted Mr. F.G. Brown, B. Sc., 
Conservator of Forests here, and he has given me 
information regarding reserves." 

"At present very little is known regarding 
species and their localities in this country and 
there is little danger of exterminating any of them. 
Rafflesia can only be protected by keeping the 
exact localities concealed and by education regard- 
ing the imaginary character of its advantages as 
a medicine in childbirth (suppose they are imag- 
inary). It grows in a forest reserve. About 
one-fifth of this country consists of forest reserves 
where normally no trees can be felled although 
the department may destroy useless species. It 
could no doubt give and enforce, to a reasonable 
extent, orders for the protection of any species 
of tree or plant found peculiar to a reserve and 
not as in the case of Rafflesia of a special value 
to the local people." 

"Several species of Conifers grow on the sum- 
mit of Mt. Poi and according to Beccari on that 
of Matang, and he mentions other interesting 
species on Matang. Mt. Poi is a forest reserve 
and Matang is kept very largely in its natural 
state by the Water Board as Kuching obtains 
her supplies from there." 

"Hunting and the collection of forest products, 
gums, fruits, etc. is permitted in the reserves; and 

the fauna, especially the birds in them and else- 
where, are in considerable need of protection." 

"The Governor of Sarawak, Sir Anthony Abell, 
is encouraging a project to make Santubong 
Peninsula a national park, also a small lake in 
the north of the country. The matter is, I believe, 
well under way." 

"When and if we have real information regard- 
ing a rare species with a limited range I consider 
that now is the time to arrange to protect it 
because at present it would be no hardship to 
anyone to avoid felling, digging up or building 
on a particular strip of ground; others equally 
suitable abound except possibly for air-strips; 
later on, if the country develops, it might seriously 
interfere with work or planning to protect some 
small locality near an industrial or agricultural 


(F. RAPPARD, Head of Forestry Service, 

Hollandia-Haven, in letter to DR. VAN STEENIS, 

October 26, 1954). Summary in English. 

"1 have discussed your letter on 'protection of 
rare plants in Netherlands New Guinea' with (my 
assistant) Versteegh and (Professor) Lam. We 
haven't at present the slightest idea what are 
rare species, nor shall we know for a couple of 
decades, and if we try to do something it will 
remain a useless project. The only way to get 
some results will be to create some reserves 
where lumbering is forbidden, but such areas 
will be in the high country. Proposals have 
already been made, also for establishing one 
coastal reserve." 


(j.s. WOMERSLEY, Chief of Division of 
Botany, Dept. of Forests, Lae, in letter to 

DR. VAN STEhNIS, August 28, 1956). 

"Firstly, I would agree with you that the funda- 
mental definition of 'rare' in terms of plant species, 
at least as far as New Guinea is concerned, will 
prove exceedingly difficult to determine for the 
practical work of this Committee (C. of Pacific 
Botany). In fact, 1 would hesitate to say that 
anyone species in New Guinea today is so rare 
that a real possibility of its extinction exists. 
As an example of this I could quote Dendrobium 
johnsonii, which immediately after the war was 
believed to be an exceedingly rare orchid which 
was faced with extinction due to considerable 
export of plants made immediately prior to 1940. 



However, further investigation has shown that 
this orchid is as common as any almost through- 
out the whole island within its own altitudinal 

"I am more concerned at this stage, at least 
for New Guinea, in the establishment of national 
parks of considerable size which could provide 
refuges against the pressing tide of agricultural 
development for the unique flora and fauna. 
The same problem has concerned the Animal 
Ecologist, Mr. K. Slater, in considering how the 
rare species of Birds of paradise may be protected 
against agricultural development of the land. 
Of course the very terrain of New Guinea ensures 
the protection of the flora and fauna although 
serious disturbances can be caused even to high 
mountain forest if agricultural development 
interferes with water-sheds and streams. " 

'There is also, at least in the Trust Territory, 
the problem of land ownership which is quite 
clearly vested with the indigenous people unless 
land is (a) purchased by the Administration, or 
(b) declared waste and ownerless. Virtually all 
land purchases to date have been designed to 
provide agricultural or residential land and only 
very few areas have been proclaimed waste and 
ownerless. This position appears to me to ensure, 
at least for several generations, that much of the 
existing forested areas quite unsuitcd to native 
agriculture will remain more or less in their 
present state." 

"However, I cannot concede that this removes 
the necessity to find some means of proclaiming 
flora and fauna reserves to protect the wild life 
of the Territory." 

"In answering your specific questions I feel the 
project is very real particularly in those countries 
of Malaysia where the population density is so 
great that there is real danger of rare species 
becoming extinct." 

"To your second question I shall be happy to 
join you as a collaborator with the proviso that 
at this stage and with our relatively limited knowl- 
edge of the distribution of plants in New Guinea, 
I would not be prepared to assert that any partic- 
ular species was so rare that it needed protection. 
We already have embodied in our Customs Ordi- 
nance adequate power to limit the commercial 
exploitation of our indigenous flora, e.g. by collect- 
ing of live orchids. Specimens of flora, fauna and 
minerals are prohibited exports except by consent 
of the administrator or his nominated delegate." 

(MR. c. A. GARDNER, in letter to DR. COSTIN.) 

Regarding plants threatened with extinction. 

"1 think that we can safely say that a number 
of our species are threatened with extinction in 
the near future, and that a number are already 
extinct. Such arc plants endemic to a small 
area. I very much doubt that the following will 
be seen again: 

Philotheca ericoides (Harv.) F. Muell 
Darwin ia carnea C.A. Gardn. 
Cryptandra eriantha Diels. 
Leschenaultia hirsuta F. Muell 
Calcadenia Drummondii Benth. 

Species threatened with extinction in the near 
future include 

Marianthus ringens (Drum & Harv.) F.M. 
Casuarina fibrosa C.A. Gardn. 
Boronia capitata Benth. 
Hemigenia viscida S. Moore 
Asterolasia grandiflora (Hook.) Benth. 

squamuligera (Hook.) Benth. 
Dryandra vestita (Ripp.) Meisn. 
speciosa Meisn. 

The above are a few that come to mind. I 
will keep your request in mind, and add to this 
from time to time. With large scale operations 
in agricultural development, it is inevitable that 
many of our localized endemics must disappear." 



The following lignose species may be in danger 
of disappearing: 

Acacia peuce F. Muell 
Choristemon humilis H.B. Williamson 
Eucalyptus serrulata Blakely and Beuzeville, a 
unique type in the genus Eucalyptus 



Three Kings Islands. 

List of species in danger, according to obser- 
vations in 1951 by Mr. Baylis: 

Alectryon grandis Cheesem. (Sapindaceae, 3 or 
4 adult trees) 



Braehyglottis arboresccns W.R.B. Oliver (Com- 

Elingamita Johnsonii Baylis (perhaps a dozen 

Hebe insularis (Cheesem.) Ckn. & Allan (Scro- 

phulariaceae, recorded as spreading) 
Paratrophis Smithii Cheesem. (Moraceae, re- 
corded as spreading) 
Pittosporum Fairchildii Cheesem. (recorded as 

Pleetomirtha baylisiana W.R.B. Oliver (Anacar- 

diaceae, only the type tree known) 
Rapanea dentata W.R.B. Oliver (Myrsinaceae, 

perhaps a dozen trees) 
Tecomantha speciosa W.R.B. Oliver (Bigno- 

niaceae, only the type plant known) 

The reason for recovery was the removal of the 


(from F. R. FOSBfcRG.) 

Island floras are peculiarly vulnerable to des- 
tructive forces of many sorts, as the areas occu- 
pied by species are frequently very restricted and 
populations are often very small. As has been 
many times reiterated, most endemic island 
species of plants are in danger of extinction if 
present tendencies toward disturbance and des- 
truction of island habitats are not checked. 

In general the actual situation seems to con- 
tinue to deteriorate throughout the insular world, 
judging by what personal observations we have 
been able to make and from reports of others, 
mostly from small indications that would be 
useless to detail here. There are a few brighter 
spots in the general scene. Major gains and losses 
have been listed in a report on the conservation 
situation for Oceania included in the report of 
the Standing Committee for Pacific Conservation, 
given elsewhere in the Congress and need not 
be repeated here. 

Work progresses slowly on the preparation of 
a file of information on the most immediately 
threatened species in Oceania which could serve 
as a guide for immediate action to save some of 
these. Reliable and up-to-date information is 
not easy to obtain, however. Contributions of 
such information are earnestly solicited. 



"The distribution of the plants considered in 
this report is that given by Jepson in his Manual 

of the Flowering Plants of California. As a 
by-product of the analysis of that work for the 
present report, I published a Tabulation of Cali- 
fornia endemics' (Leaflets West. Bot. 7, 1955), in 
which I discuss matters relating to this important 
and distinctive feature of the California flora. 

Genera endemic in California. 

(a) Known to occur in a State or National 

m Scquoiadendron Buchholz (Taxodiaceae) 

Sedella Britt. & Rose (Crassulaceac) 
m Draperia Torrey (Hydrophyllaceae) 
m Phalaeroseris Gray (Compositae) 
m Orochaenadis Cov. (Compositae) 
m Gilmania Cov. (Polygonaceae) 

Oreonana Jepson (Umbelliferae) 
m Whitneya Gray (Compositae) 
m Holozonia Greene (Compositae) 

To these will perhaps have to be added the 
following which probably can be found in a pre- 
serve : 

m Heterogaura Rothrock (Onagraceae) 
Pseudohahia Rydb. (Compositae) 

(b) Not known in a Park or other Preserve, 
"National Forests" not considered: 

m Etosperma Swallen (Gramineae) 
m Odontostomum Torr. (Liliaceae) 
m Carpenteria Torr. (Saxifragaceae) 

Acanthomintha Gray (Labiatae) 
m Holocarpha Greene (Compositae) 
m Crockeria Greene (Compositae) 
m Tracyina Blake (Compositae) 
m Neostapfia Davy (Gramineae) 
m Hollisteria S. Wats. (Polygonaceae) 
m Lyonothamnus Gray (Rosaceae) 

Monolopia DC. (Compositae) 
m Blepharozonia Greene (Compositae) 
m Eastwoodia Brandagee (Compositae) 

Genera preceded by an m are monotypical. 

The following trees, endemic in California, 
are found in a State or National Park: 

Abies magnified Murr 
Cupressus macrocarpa Hartw. 

Sargentii Jepson 
Pinus balfouriana Jeffrey 

,, monophylla Torrey 

muricata Don. 

ponder osa Dougl. var. Jeffreyi Vasey 

sabiniana Dougl. 



Finns torreyana Parry 

tuberculata Gord. 

Scquoiadendron gigantewn (Ldl.) Buchholz 
Torreya californica Torr. 
Lithocarpus densiflora (Hook.&Arn.) Rehd. 
Quercus chrysolepis Liebm. 

Douglasii Hook. & Arn. 
lobata Nee 
morehus Kellogg 
Platanus racemosa Nutt. 
Aesculus californica (Spach) Nutt. 
Fraxinus dipclala Hook. & Arn. 

These are not found in Parks or other 
Preserves : 

Abies bracteata (Dougl.) Nutt. 
Cupressus Forbesii Jepson 
go vcniana G ord . 
inacnahiana M u r r . 

var. Baker i Jepson 
nevadensis Abrams 
pygmaea Sargent 
Sargent ii var. Duttonii Jeps. 
Pinm contorta Dougl. v. Bolanderi Vasey 

,, Coulteri Don. 
Pseudotsuga macrocarpa Mayr. 
Taxus rev if olid Nutt. 
Quercus dumosa Nutt. var. Macdonaldii Jeps. 

Wislizenii A. DC. 
Jug Ian s californica Wats. 

Hinds ii Jepson 
Lyonothamnus calif or nicus Gray 
Acer negundo L. var. californicum Sargent 

The California Floral Province, extending from 
southern Oregon to northern Baja California, 
with exclusion of considerble areas in the eastern 
part of the State related floristically to the Colum- 
bia Plateau, Great Basin or Sonoran Desert, was 
proposed by me in 1957 (Leaflets West. Bot. 8:5). 
The number of endemic genera is 65." 


National Park of Talinay, Prov. of Coquimbo. 

Visited on April 30, 1955, by Dr. and Mrs. 
Skottsberg, accompanied by Mr. Carlos Jiles, 
a resident of Ovalle. The forest was found in 
primeval condition and in no way threatened ; it is, 
in fact, in better condition than the nearby quite 
similar Frai Jorge forest north of the mouth of 
Limari River and has been much less visited than 
this. The latter has been well surveyed by Messrs. 
Munoz and Pisano, and before by Dr. and Mrs. 
Skottsberg (publication in Acta Horti Gotoburg 


18, 1950). A corresponding survey of Talinay 
ought to be undertaken. The composition is the 
same in the two places, but Cerro Talinay is about 
30 m. higher (700 m.) and makes an impression 
of being even more humid; both are enveloped 
in fog and the vegetation drips with water. Nei- 
ther is of difficult access; in the case of Talinay 
a motor road leads to the foot of the mountain, 
and from there the ascent is made on foot through 
dense chaparral, where, in places, Puya chilensis 
is dominant. The forests are famous as consist- 
ing of a highly hygrophilous community, sur- 
rounded on all sides by xerophytic vegetation and 
separated from the south Chilean rain forests 
by many degrees of latitude. The great Pan- 
american Highway will pass one km. from Tali- 
nay, and it is expected that the peculiar forest will 
be of great touristic value. It should be remem- 
bered that the soil cover of delicate ombrophilous 
herbs and bryophytes, where the foot sinks 
deep, cannot bear any traffic at all without 
getting damaged and that the extreme rain forest 
type with lianas and epiphytes covers only a 
narrow strip. The distance from the Highway 
to Frai Jorge is about 20 km. 


In my Report for 1933-38 to the Sixth Pacific 
Congress I told of the establishment, in 1935, of 
the Juan Fernandez National Park, the first of 
its kind in Chile. Even if, as little was done to 
enforce the regulations, an efficient conservation 
of the native vegetation still remaining wasn't 
in any way ensured, nothing like the devastation 
that has taken place in the last twenty years could 
be expected. That much would have happened 
between 1917, when I surveyed the islands, and 
1935, could be foreseen; the intentional introduc- 
tion to Masatierra of one of the dangerous pests 
from the mainland, a European species of Rubus 
(so far determined ulmifolius) had been reported, 
and the renewal of Masafuera as a convict settle- 
ment in 1927 ought to have left its marks. In 
order to find out what had happened and to 
make a comparison between 1917 and now, I 
returned to the islands in December, 1954, and 
spent three months there. Also this time I was 
accompanied by Mrs. Skottsberg. 

Masatierra. The settlement remains a fishing 
village, but it had grown considerably, from 
about 200 to about 600 people, which is too much, 
as too many persons are not engaged in the 
fishing industry. As before, the population 
concentrates around Cumberland Bay, the only 



harbor. The lack of suitable land and the exceed- 
ingly broken topography forbids agriculture if 
not on a miniature scale, and horticulture showed 
little increase, even if small cultivations had 
extended into the valleys east of the Colonial 
Valley, but this had little to do with the destruc- 
tion during the thirty-eight years that had passed 
since our previous survey. 1 do not think that 
the number of cattle, about 250, was much 
greater than before; the difference was that there 
was almost no pasture left, and we shall see the 
reason. Sheep had been introduced and allowed 
to increase without any restriction at all. The 
whole island lay open to them; they had invaded 
the drier forest type and the brushwood of the 
high ridges; through the action of a number of 
enterprising individuals the national park had 
been turned into a sheep farm. The official 
figures stated about 3,750 sheep, of which 3,100 
grazed the lower treeless western part and the 
remainder the denuded slopes of the easternmost 
valley of Frances, but in addition numerous 
others were observed also in other valleys and 
on the high ridges so that the total number must 
have exceeded 4,000. Certain areas had been 
turned into a desert, and the valley systems of 
Frances, Ingles, and Villagra offered a horrible 
sight. Once the soil formed by the easily disin- 
tegrating basalt, had been deprived of its plant 
cover, native (as in Villagra) or not, erosion set in 
with full force. Horses are of slight importance 
as communication between the coves is so much 
easier by motor boat; the old trails to Villagra 
and Frances are seriously damaged and almost 
dangerous; the lower part of Villagra, in 1917 
covered by native grassland, had changed beyond 
recognition, and the zig-zag trail across Salsi- 
puedes to Ingles valley, worn deep into the rock, 
had been completely effaced. 

Goats, introduced in the 16th century, had 
become naturalized and must have been very 
plentiful once, but in the 19th century their 
number had become so much reduced that they 
were protected for sentimental reasons because 
they descended from the goats of Alexander 
Selkirk, with whom Defoe's Robinson Crusoe was 
identified. In 1916-17 few were shot because 
they had retreated to places of extremely difficult 
access, and in 1955 they were scarce and not 
seen by us. But there was good help for this; 
under the pretext of providing goat's milk a 
small herd of angora goats had been imported 
to the colony, where they took to the mountains 
at once, multiplied and spread. One day 1 counted 
sixty on one of the steep ridges above the settle- 
ment; being pure white, they are easily discern- 

able even from a large distance, and some were 
seen later on the most inaccessible precipices. 
They are private property so that hunting them 
is illegal. Two more additions to the fauna should 
be mentioned, Nasua rufa, introduced to help 
combat the big rats, and rabbits. Nasua prefers 
the dense forest and is said to be scarce; it is an 
excellent climber and accused of liking birds' 

It is hardly probable that, in old times, much 
damage was done in the native forest after the 
wild pigs had been exterminated, except where 
logging was practised, until the arrival of the 
terrible Chilean pest, the shrub Aristotelia 
chilemis (maqui) some time before 1854. In 
1917, it filled all the valley bottoms and lower 
slopes above the everywhere denuded coastal 
belt, penetrating the native forest and prevent- 
ing, by its deep shadow, the endemic trees all 
native trees are endemic from germinating, so 
that their disappearance from the lower forest 
belt was foreseen already by Johow in the 90's. 
The maqui had continued its progress and ad- 
vanced into the upper forest belt, reaching the 
ridges in 500-600 m. in many places; the fruits 
are dispersed by the thrush. Another Chilean 
shrub, Ugni Molinae, locally known as murtilla, 
introduced on purpose because of the aromatic 
berries during the latter half of the 19th century, 
was, in 1917, practically confined to the barren 
slopes above the Colony. It had increased tre- 
mendously and invaded the native shrub along 
the steep ridges, the home of many of the rare 
endemic genera and species of plants, some of 
them now on the verge of extinction. All our 
good old collecting grounds were more or less 
ruined. A special list of rare and vanishing 
species has been prepared. 

One of the most precious and scientifically 
interesting forest products, the endemic and 
monotypical palm Juania australis, since long 
protected by law, has almost entirely disappeared 
from the accessible parts of the forest belt, but 
freshly manufactured curios of it were still 
offered for sale. 

The Pangal Canyon, the only typical canyon 
in Masatierra and a gem, will serve as an example 
illustrating the changes since 1917. Toward the 
sea was a fine stand of Boehmeria excelsa along 
the stream, and the interior had fine native 
forest with stately tree-ferns right up to the ter- 
minus, a verical cliff wall with a cascade. Of all 
this vegetation, nothing remains; the outer slopes 
are naked, full of weeds and grazed by cattle; 
the interior, a jungle of maqui; the waterfall is 
reached using the tunnels made by the cows. 



The bramble-berry, introduced in the 20's 
by a colonist living in the Anson valley, to be 
used as a living fence around his orchard, had 
destroyed it and spread like wild-fire up the valley; 
the small open place at the foot of Mt. Yunque, 
often visited by us and surrounded by fine forest, 
was now changed into an impenetrable thicket. 
From here it spreads by man and birds the fruit 
being edible in all directions and pioneer speci- 
mens florish on the high ridges above 500 m. If 
nothing is done to stop it, it will invade the island; 
the people complains of the "zarzamora," but 
nothing has been done. It may be too late now 
and will, in any case, cost a lot of money and labor, 
a matter for the high authorities on the mainland. 

It goes without saying that the changes also 
affect the fauna. The endemic land birds are 
decreasing, and the peculiar pacific land shells 
will not, I presume, find the maqui a suitable 
biotope. The insect life, mostly small incon- 
spicuous forms, is still surprisingly rich, at least 
on the higher forested ridges; numerous genera 
and perhaps some 90",, of the species are endemic. 

Whether fires have played any important role 
I cannot tell, but I was told that in 1930, a fire 
swept Ingles valley, and very likely this is true. 
It is, anyhow, ruined; the pasture is gone, and 
sheep and cattle left to starve to death. They 
cannot get out. 

Santa Clara Island, a satellite of Masatierra, has, 
as far as we know, never borne any forest growth, 
but was once covered by grassland sprinkled with 
a few dwarf rosette trees belonging to the endemic 
genera Rea and Dendroseris. All this is gone with 
the exception of some specimens growing in inac- 
cessible gorges. The islet is overstocked with sheep 
and looks like a desert; in 1955 the sheep num- 
bered about 500, but we saw no goats, once said 
to be numerous "Goats Island" is an old name 
for the place. The detached rock Morro del 
Spartan, separated from the islet by a narrow 
channel not crossed by sheep or goats, gives some 
idea of what the scenery must have been in bygone 
time. This morro is the type and only locality for 
Chenopodium Sanctae Clarae, of which half a 
dozen specimens were observed. 

All visitors agree that Santa Clara lacks fresh- 
water, but rain falls during the winter months 
when some water must be found, otherwise 
sheep could not exist. According to recent 
information the owner of the sheep was going to 
evacuate the island. 

Masafuera. The law says that no permanent 
habitation shall be set up on this island ; and when, 


in 1930, the convict station was abandoned, 
Masafuera was left uninhabited. However, in 
1940, a fishing colony was established at the 
entrance to Casas valley and took possession of 
the government buildings. Langust fishing, 
the principal industry of Juan Fernandez, is 
profitable around this island, and the colony, 
though living under rather primitive conditions, 
was prosperous. No cultivations of any kind 
exist now, there were only about thirty head of 
tame cattle, a few horses, and twenty sheep, so 
that the damage done was negligible. In 1917, 
the marks of the first penal settlement were very 
conspicuous; the accessible forest patches had 
been logged and grazed; huts had been built in 
the uplands; potatoes and vegetables were 
grown, but the high land south of the Vacas 
valley, crowned by the summit, 1,570 m. above 
sea level and extremely wet, was covered by an 
impenetrable fern forest and showed almost 
no sign of human activity. In 1917, good forest 
groves were found in the upper gorges of the 
valleys all along the east side of the island the 
western side is a precipice between 300 and 
700 m. above sea level, the maqui was rare, and 
the grassland higher up, with its extensive fields of 
big ferns and bands of Gunnera along the streams 
and scattered stout specimens of Dicksonia, 
made the impression of a peculiar fern savanna 
above 900 m. Goats were very plentiful, and many 
plant species confined to places they could not 
reach. And now, in 1955, this wonderful island 
offered a very sad sight. The forests had been 
logged and were full of maqui; the first bram- 
bleberry shrubs had made their appearance. We 
had difficulty in recognizing places we used to 
know so well and now found utterly ruined, a 
result of a series of devastating grass and bush 
fires. The dates given were 1939, 1942 or 1943, 
and 1944, and one of them, which swept the In- 
ocentcs ridge to the highest summit of the island, 
was not accidental. Another fire had run over 
the Barril ridge across to the gorges on the 
western side. The fern savanna was gone; 
charred Dicksonia trunks told the story. Forest 
groves of small size in the hanging gorges cut 
into the west wall of the island are still unharmed. 
The only large patch of closed wood is at the 
northern end. This was visited. It is of a dry 
type on the flat ridge between two valleys. In 
this part, natural Stipa grassland, very monoto- 
nous, covers extensive areas; otherwise the up- 
land is overrun by Anthoxanthum odoratum and 
Rumex acetosella, just as it was in 1917, and the 
native Alpine species are scarce except close to 
the edge of the west wall in 1,100-1,400 m. 





Botanic Gardens, Bogor, Indonesia. 

If we may compare botany to a tree, we can say 
that it has its roots in Europe but that it is coming 
into flower in the tropics. Most of its flowers, 
however, are still in bud. To cause these buds 
to open is, in brief, the task of botanical institutes 
in the tropics. We mean: to realise the splendid 
possibilities for scientific research provided by 
the wealth of tropical forms. 

With respect to the taxonomic study of a trop- 
ical flora, the task of an institute in the tropics 
is more complicated than that of an institute in 
a temperate climate. The latter can be merely 
a "working" herbarium where botanists deal with 
the material that comes to their hands. But a 
tropical institute not only has to elaborate 
material for the purpose of compiling taxonomic 
publications, it also must supply the material 
itself, by collecting it on expeditions in the field, 
an undertaking that may be too expensive or too 
complicated for an institute in a temperate 

In the course of history, every botanical insti- 
tution has had its ups and downs, but this must 
not divert our attention from distinguishing the 
two kinds of herbaria: the "working" herbaria 
and the "supplying" herbaria. An example of 
a working herbarium is the Kew Herbarium. 
Here botanists are preparing and publishing 
revisions, but their material must come from 
somewhere else. An example of a supplying 
herbarium is the one at Sandakan. Here no 
proper taxonomic work is performed, but material 
is collected for the Kew botanists and others. 
The Bogor Herbarium has always both produced 
taxonomic studies and supplied material ; the same 
is true for the herbaria at Singapore and Manila. 

A modern taxonomic publication, especially 
if the author aims at a high standard, is the result 
of the cooperation of many people. Specialists 
are spread all over the world; the type material 
of Malaysian plants is scattered over at least 
a dozen herbaria, and for good results so much 
collaboration is necessary that we sometimes 
wonder whether the task of the Keeper of a 
modern herbarium is anything else than shipping 
and inserting material and sending botanists 
abroad. In any case, if we wish not to botanize 
alone but to contribute to international botany, 

our best policy is to show how we can be most 
useful to others. This can be done by utilizing 
the specific advantages of the site of our institute. 

We need people who are able to collect critically 
and to make notes in English to go with the 
material, thus giving important information to 
the herbarium taxonomist. The collector must 
be prepared to take at least three duplicates a 
dozen or more is still better. Further, it is ex- 
tremely useful for him to have so much knowledge 
of the existing forms that he can pre-identify the 
material collected. At Bogor, Mr. Nedi and 
Mr. Noerta perform this work. Although lacking 
advanced schooling, they have accumulated exten- 
sive knowledge during field work with botanists 
and continued training themselves in recognizing 
families and genera. They are invaluable because 
in this respect they have achieved what a pro- 
fessional botanist achieves only after long, long 
experience. Therefore, if a man is found who 
shows talent in recognizing plants, he should be 
taught and encouraged, no matter what his status 
is. Even a common labourer could thus become 
one of the treasures of the institute. 

Finally, duplicates of the material collected 
must be distributed. This is by no means easy. 
The material must be divided into adequate 
portions and labels must be copied carefully. 
All this material must be packed and shipped. 
The dispatch of duplicates is not one of the minor 
occupations of the Bogor Herbarium staff; we 
need mention only that in 1956 we shipped a 
total of seventy-four packing cases of duplicate 

A herbarium connected with an important 
tropical garden has another special function to 
fulfill, namely to collect and depict garden 
material and to give information asked for. A 
tropical garden containing many "wild" species 
can provide more direct knowledge than a garden 
in a temperate climate. 

Because the functions of a tropical garden are, 
as we have seen, quite varied, it can easily be 
understood that we must prepare ourselves not 
to lose our heads when we have to face difficulties 
resulting from lack of funds or lack of trained 
staff. The most important thing is to keep what 
we have. We therefore consider the alcohol and 



the corrosive sublimate that preserve our speci- 
mens to be the life blood of the tropical herba- 
rium. Almost equally important are the tins and 
the paper that protect them against breakage 
and dust. 

It is similarly important to improve our 
material. When material required for a revision 
is sent out on loan and later returned and inserted, 
the new identifications by specialists will help us 
in turn to identify new specimens by comparison. 

As apparent from the above, a well-trained 
technical stafT is of the utmost importance. A 
herbarium with only a botanist is nothing. A 
herbarium with good technical staff is at least 
something. This contradiction holds good 
especially in the tropics. 

The work of a botanist will stimulate other 
activities: collecting and distributing duplicates 
and gaining information from living material. 
Of course he should work and publish, but he 

must remain aware of the fact that an excellently 
prepared and annotated plant specimen, widely 
distributed, is better than a second-rate publi- 

With respect to the creation of new local 
institutions, it is well to recall the function peculiar 
to the tropical herbarium, namely to supply 
material to the great international body of 
science. A library as extensive as that at Bogor 
is necessary to operate a "working" herbarium, 
and it must be kept up to date. But a properly 
functioning "supplying" herbarium requires only 
an active botanist with a few willing local men, 
the necessary collecting and drying equipment, 
and facilities for shipping the material. 

The tree of botany has become too large to be 
tended by one institute only. For efficient growth, 
proper flowering, and plentiful fruit, many 
gardeners are necessary, each endeavouring to be 
the right man in the right place. 


C.G.G.J. VAN STEENIS: I congratulate Prof. Kusnoto 
on his excellent exposition of the functions of tropical 
herbaria. I am glad to learn that the Bogor institution 
intends to combine supply of material to other herbaria 
with work on the flora, as indeed all tropical institutes 

S.Y. HU: One must be careful to collect and distribute 
enough duplicates of each specimen. 

p.s. ASHION: May not too many specimens for identi- 

fication or as representatives of local collections embarrass 
the major herbaria and strain their facilities? 

r.G.G.j. VAN SIEENIS: Well annotated, fertile material 
will always be gratefully received by the herbaria in the 
temperate regions. 

H.M. BURKILL: The large herbaria still need such 
material, even of common species. Duplicate specimens 
are the best currency available to botanical institutions. 





Botanist of the Forest Research Institute* Bogor, Indonesia. 

The problems discussed here deal with Botanic 
Gardens and Herbaria. Those problems may be 
divided into three categories: Time, Climate, 
and Man. 

Botanical investigation in its initial state was 
taxonomical and started outside the tropics, 
although its aim was the study of tropical plants. 
For centuries, materials, which served as bases for 
descriptions, floras, etc., have been heaped up, 
first in European and later also in American 
herbaria. Tropical institutes, like the Botanic 
Gardens in Bogor, the Calcutta Botanical Garden 
(in a lesser way), the Rio dc Janeiro Garden, 
were more or less considered transit-institutes to 
facilitate collections to be sent abroad. As soon 
as foreign botanists withdrew from such institutes, 
the work came usually more or less to a standstill. 

The reasons are obvious: tropical institutes are 
usually situated in countries where science is in 
its initial stage or the local population has not yet 
reached the stage of development for pure scien- 
tific research. These comparatively young nations 
have other, more important problems, which 
confront them. The repercussions of a weak 
economy were felt several times during the English 
and Dutch Colonial period; in poor times, the 
Bogor Institute was deprived of its staff, and work 
came to a standstill, while the collections were 

One of the main difficulties, confronting the 
Herbaria of young countries, is that most of the 
type material is abroad and not represented in 
the country itself. This is a big handicap, as it is 
a time and money consuming work to study such 
specimens abroad, as several European Herbaria 
are unwilling to send material on loan for fear of 
damage and loss (which is quite understandable). 
On the other hand, we should never forget, that 
because collections were kept outside the tropics, 
many of the older collections are still in good 
condition. Without these collections, progress 
in taxonomy would be much hampered. 

The young tropical countries, which will take 
up now the completion of taxonomic and floristic 
work are confronted with almost insurmountable 
difficulties. We will mention only a few. Taxo- 
nomy in Europe and America is not very much 
appreciated by the "man in the street," who 

t Presented by Kusnoto Setyodiwiryo. 

considers it a waste of time and money and hence 
these governments pay poor salaries for too few 
jobs, and funds for working arc scarce. As these 
conditions prevail in European countries, we can 
imagine, that they are far worse in tropical 

There is only one remedy: education. The 
wealth of the vegetation in the tropics can never 
be made useful to man without thorough investi- 
gation, which must be carried out in botanical 
institutes and must start with inventory, which 
means taxonomic work. 

Besides lack of funds, there is a lack of qualified 
botanists. The jobs arc not attractive, as payment 
is poor and the study long and difficult. For a 
taxonomic botanist, knowledge of four or five 
foreign languages is an absolute must, which 
means that a student in Indonesia, who is taught 
only one foreign language in secondary school, 
has to spend much time and energy to make up 
for the other languages, and this is often a hope- 
less task. 

Then comes problem of maintaining existant 
herbaria and collections of new material. 

It is certainly not true that in the humid tropics 
the dried material deteriorates very quickly. That 
depends on how the material has been preserved 
and how it is stored. Attack by insects is suc- 
cessfully checked by poisoning the specimens 
with an alcoholic solution of corrosive sublimate. 
Bulky parts of specimens, like fruit, where the 
sublimate does not penetrate, are likely to be 
attacked and part of this material should be 
conserved in spirits. The glueing of the specimen 
on the mounting sheet by means of a poisoned 
glue prevents further attack of insects. We could 
observe that the material in the Singapore Herba- 
ria was far less attacked than that in Bogor; in 
Bogor the material was not glued on (nowadays 
the method of glueing has been adopted too). 
The glueing does not interfere much with taxo- 
nomic investigation, as tepid water easily loosens 
the parts, which can be taken off for investigation. 
On the other hand, the advantages of glueing are 
enormous. Not only is the deterioration of the 
plant checked, not only is insect damage less, but 
the damage caused by handling is reduced to a 



minimum. The damage done by handling and 
dispatching the Bogor material has done so much 
damage that the material must be considered 
poor in comparison with that of Singapore. 

As herbaria in the tropics are usually open 
buildings, infection is more likely than in colder 
climates. This may be prevented by storing in 
tin boxes with tight fitting lids; an insect repellant 
might help to chase insects. 

There is, however, a great danger for specimens 
on loan from abroad which are often not 
poisoned. To young workers this is often not 
known, and the stored material may be finished 
by insects in a couple of weeks. Storage of such 
material is only possible in hermetically sealed 
boxes, provided with an insecticide. As the 
material, however, is handled, this is not much 
good. The safest way, which we have adopted 
in Bogor, is to poison the loaned specimens in 
a pure alcoholic corrosive sublimate solution, 
wherein the mounted sheet is immersed for a short 
period. The sheet remains clean, and the poison- 
ing is invisible. 

Upkeep of a collection provided the funds 
are there docs not add any problems, as this 
routine work is within reach of a lower-paid 
official. In Bogor, however, we have trouble in 
filing the material properly. Personnel doing this 
have practically no inkling of scientific names, 
and mistakes are numerous. Again, this is a 
problem which could easily be eliminated by a 
better paid official with a higher education. 

Next comes the collecting of new material. 
If the herbarium in a tropical country wants to 
stand on its own legs, it should collect as much 
material as possible to replace type specimens as 
far as possible. A condition, as prevailing in 
Bogor, where only half or less of the known and 
described plants are represented is an unsatis- 
factory condition. 

Collecting, however, is possible only by means 
of a team of qualified botanists with ample funds. 
So long as these two requisites are not fulfilled, 
the work should be postponed, as we have learned 
from experience in the Forest Research Institute 
that indiscriminate collecting is a waste of time 
and money. 

A "Botanical Survey/' as set up by the Indian 
Government to initiate and guide an over-all 
survey, has much to commend itself, provided it 
has enough funds and qualified personnel (which 

is for the time being not the case in Indonesia). 

As matters stand in Indonesia, where no flora 
is extant and most of the type material is not 
available in the country, the only alternative was 
to continue the taxonomic survey in the way it 
has been done for almost 200 years. A body has 
been created, the Flora Malesiana Foundation, 
initiated and under the able directorship of 
C.G.G.J. Van Steenis in Leiden and Kusnoto 
Setyodiwiryo in Bogor, to publish the flora of 
not only the area of Indonesia, but also the Phi- 
lippines and Malaya. The bulk of the scientific 
work is done in Leiden, where a botanical staff 
has at its disposition an adequate library and 
necessary herbarium material. The funds for 
this important work are provided by the Indone- 
sian Government. Moreover, the Bogor Botanic 
Gardens take an active part by providing herbar- 
ium material. 

It is in this way only, that botanical investiga- 
tion may be carried on in Indonesia. In the 
future, Indonesian taxonomists should take an 
active part in this large scale important work. 
For the time being, Indonesia may contribute by 
increasing its activity in collecting material all 
over the area and in providing the means (funds 
and help) to support the foreign taxonomists, 
who work on this flora on a non-profit base. 

Again it is proved here that more than any 
other science taxonomy is international, and the 
whole world benefits from the scientific cooper- 
ation of nations. 

The two main problems of the Botanic Gardens 
in Bogor are lack of funds and (as is the case 
with numerous, also extra-tropical gardens) a 
difficulty in getting the collections properly 
named. For a large garden as that of Bogor 
which consists mainly of woody plants (amongst 
which are several undescribed ones), a qualified 
taxonomist is urgently needed. If a professional 
botanist is added to the Garden Staff, the danger 
that the importance of the collection will decline 
is eliminated. With a non-botanical staff, the 
trend is more to raise ornamentals, and the scien- 
tific collection may be neglected. 

Continuous collecting of seeds and seedlings 
of botanical, more than of ornamental or business 
interest, is an absolute must. This is, even under 
prevailing circumstances, always possible, pro- 
vided the staff is convinced of the scientific im- 
portance of the Garden. 


C.G.G.J. VAN STLENIS: I consider the material at Singa- 
pore and Bogor to be of similar quality. Glueing specimens 


to sheets was developed to reduce accidental loss (or theft) 
of portions of the specimens, but this practice is most 


troublesome to research botanists. Despite the greater o. SEIDENFADAN: Taxonomic research is the essential 

cost in paper and space, wrapping each specimen in a basis for economic development of natural resources; 
separate cover is the best protection against friction damage governments must realize this and give adequate financial 
and loss ot detached fragments. support. Institutes must correct the opinion that such 


P.R. wvrntRLhY: 8-hydroxy-quinoline-potassium sul- r.c,.j. VAN SIEENIS: We should all endorse His 

phate may be used as a substitute for, or in addition to, bxcellency's statement, may we record this? (There were 

mercuric chloride. Paradichlorobenzene is a valuable insec- no dissenting voices, and the meeting appeared to be in 

ticide in herbarium cabinets. general agreement.) 





Professor oj Botany, Impci ial College of Tropical Agriculture, Trinidad. '\ 

This paper is a precis of PURSEGLOVE: 
History and Functions of Botanic Gardens with 
special reference to Singapore: Tropical Agri- 
culture 34(3), July 1957. 

Botanic Gardens are places at which to study 
plants. The earliest were herb gardens, the first 
being that at Pisa founded in 1543. From then 
through the Linnean era to the present day, 
gardens have developed on scientific concepts, 
accumulating not only living plants but herba- 
rium material and literature, some on a world- 
wide basis, some on a local basis. 

Though many gardens were established in the 
16th and 17th centuries in Europe, the earliest 
botanic garden in the tropics is thought to have 
been that at Pamplemousses in Mauritius, es- 
tablished in 1735. In that century and the next, 
many tropical gardens were set up, chiefly with 
the object of studying plants of economic impor- 
tance either local or introduced. In Asia, the 
gardens at Calcutta were founded in 1786; at 
Penang (the first of three), about 1796; Buiten- 
zorg (Bogor), 1817; Peradeniya, 1821; and at 
Singapore (the first of three), in 1822. These 
gardens and the men who have served in them 
have laid the foundation for the scientific study of 
Asian botany and of much of the forest and agri- 
cultural economy of the tropics. One has only 
to consider the tremendous value to the world 
the following tropical agricultural plants have 
and peculiarly that most of them were removed 
from their natural habitat to alien lands: Hevea, 
cacao, quinine, coffee, cloves, nutmegs, sugar, 
bananas, limes, vanilla, cassava, sweet potatoes, 
maize, and many others. 

In Malaya, the first Penang Gardens were 
founded soon after the establishment of a trading 
station by the East India Company in 1786. 
Similarly the first gardens in Singapore were 
founded but three years after the trading post 
was set up. The attention of both was given to 
the growing of nutmegs, cloves, and other crops 
of commercial value. Of the surviving successors 
to these Gardens, that at Penang is now a pleasure 
gardens; and only that at Singapore is a botanic 
garden in a true sense. This latter came into being 

t Presented by H.M. Burkill. 

t Formerly Director of Botanic Garden, Singapore. 


in 1859 as the garden of an Agri-horticultural 
Association. Within a few years, the government 
had taken over its administration; and its work 
expanded to include a study of the flora of the 
Malay Peninsula, as well as work on crops of 
possible economic value, such as Cinchona, 
coffee, eucalyptus, ipecacuanha, tea, maize, 
sugar-cane, colanuts, mahogany, oil palm, cacao, 
and, of course, the classic pioneering of Hevea. 

The Singapore Gardens made the greatest 
progress in laying the foundation for agriculture 
and forestry in Malaya under Ridley in the two 
decades following 1888 when he arrived as 
Director. Ridley also conducted the most exten- 
sive research into the flora of the country. He 
founded The Agricultural Bulletin of (he Malay 
Peninsula in 1891 to be superceded by The Agri- 
cultural Bulletin of the Straits and Federated 
Malay States in 1901. 

Through the researches of the Botanic Gardens, 
the Departments of Forestry and of Agriculture 
were set up to extend the work of their respective 
lines. This did not mean that the Botanic Gar- 
dens ceased their interest in these matters. The 
work of all generations of members of the Depart- 
ment has had a bearing on the evaluation of 
Malaya's inherent wealth in its vegetation and 
in its agri-horticultural potential which is now 
made known through its serial publication, The 
Gardens' Bulletin, and The Revised Flora of 
Malaya. Even during the Japanese interregnum 
in Malaya, work at the Gardens continued under 
the direction of Japanese botanists. Though this 
period was fraught with difficulty, the herbarium 
and library suffered no loss, but the living col- 
lection of plants was sadly reduced. In the post- 
war years, the Gardens quickly regained their 
former good condition, but these latter years have 
been characterised by lack of trained staff. 

The work of a botanic garden is botanical, 
horticultural, and educational. For the first 
purpose, the garden must be a museum of living 
plants to compliment the herbarium, a museum 
of dried plants. In the Singapore Gardens, some 
3,000 perennial species grow, not counting hy- 
brids. The herbarium contains about 400,000 

sheets, mostly of Malaysian origin, whose quality 
and quantity are constantly being increased by 
loans for revision and by exchanges of duplicate 
material. Collecting is done in Malaya and in 
neighbouring countries where the vegetation has 
botanical affinities. This work leads to taxonomic 
studies which are necessary preliminaries to the 
preparation of floras. With the complexity of 
Malaysian botany, this work is a tremendous 
undertaking. Over the past one and a half 
centuries that Malaysian botany has been studied, 
an immense amount of information has been 
gathered. In 1921-24, Ridley published the first 
Flora of the Malay Peninsula. So much more 
data are now available that this work is in urgent 
need of being superceded by a revised Flora, a 
work which has already been begun. Similarly 
the Singapore Gardens is throwing in its lot 
with the immensely more difficult task in the pre- 
paration of a regional flora, The Flora Malesiana. 
It is only from such undertakings that man can 
live, for, to quote Linneus "Science, and in the 
first place botany, is the only reliable basis of 
private as well as national economy." 

On the horticultural aspect, the exchange of 
seed and the acquisition of species of possible 
value from other countries are equally important 
activities. It is the natural sequel to the botanical 
research just mentioned. From it has arisen, 
for example, the immense and vitally important 
Hevea planting industry. Besides Hevea, there 
are numerous other plants, of less apparently 
over-riding importance but nevertheless of value 
commercially (e.g. oil palm, cacao), medically 
(e.g. Rauwolfia spp.), and aesthetically (e.g. 
Mucuna, cacti, orchids). The work on agricul- 
tural plants is now the responsibility of the 
Department of Agriculture, but nevertheless the 
Gardens continues to play an important part. 
Horticulturally Malaya's relatively seasonless 
climate is a difficulty for many spectacular orna- 


mental do not receive the required stimulus to 
flower. Breeding work has been done on some 
groups of plants to select responsive varieties, 
and the main breeding work at present being 
done is on orchids. 

Educationally the Gardens help to increase 
people's, in particular children's, interest in plants. 
School parties regularly visit, and the Gardens 
supplies material to schools and the University 
for teaching and planting. A considerable 
amount of advisory work is also done and active 
encouragement is given to such bodies as the 
Singapore Gardening Society, the Malayan 
Orchid Society, and the Malayan Agri-Horticul- 
tural Association. 

In addition to the foregoing, the Gardens has 
the responsibility of running the Nature Reserves 
on Singapore Island in which all wild life is pro- 
tected. The botanical and educational aspects 
of the Gardens' work are represented in this 

The future, with political maturity of Malaya 
and Singapore, holds two cogent problems. 
Firstly, there is the question of trained staff, for 
much will depend on the new recruits who will 
have to take over the duties heretofore exercised 
by overseas men, and who it is to be hoped will 
carry on the traditions of competent research 
built up over the past century. Secondly, there 
is the question of finance, a satisfactory solution 
to which rests on the ability of the staff to demon- 
strate to politicians and the public the worthwhile 
role the Gardens plays in the economy and pres- 
tige of the country. Two and a half centuries ago 
the following lines were written about botany: 
"It is certain, however, that this Science, like all 
Sciences, flourishes sometimes more and some- 
times less, all in accord with the inclination of 
Rulers and the Favour of Government." 
(Commelijn) How true it is still today! 


p.w. RICHARDS: The collections of living plants in the 
tropical gardens must not be neglected, nor limited to 
economic or ornamental species. Visiting botanists find 
them invaluable for a wide range of studies. 

J.H. HURUMANN: Private institutions are faced with 
the same difficulties as official organizations, and so their 
utility is limited. 






Botanic Gardens, Singapore. 


In a period of transition of administrative 
authority such as is going on in Singapore at 
present, it is an advantage to maintain a sense of 
humour. Many odd things crop up, like the 
zealous Treasury official who, at an application 
for funds to buy some more herbarium cabinets, 
quoted Government's General Orders which 
require a Director of a department to review his 
department's records when they become five years 
old with a view to destroying those of no further 
import. Strictly correct, of course, but when do 
botanical records (herbarium specimens) reach 
an age for destruction ? Or again, another official 
dealing with equipment for the Gardens wanted 
to know how many persons normally sat on a 
garden seat. He had plainly yet to learn the facts 
of life that in Singapore's predominantly ado- 
lescent population, "two's company; three's 

These points will serve to show that in the 
growing political responsibilities of the new 
nations of Southeast Asia there is now a cadre of 
administrative officers with everything to learn 
in the shortest of time. Rome was riot built in a 
day. An intelligent inquisitivcness is a healthy 
sign amongst this new band of administrators, 
but the simplest way of learning is by coming to 
see and personal discussion, and one could 
wish that this, disrupting though it may at times 
be to normal routine, was more often practised. 
Though it may be adequate to copy one's pre- 
decessors, it is desirable that those who hold the 
purse-strings and those who run the department 
maintain a close liasion. Thus one may hope to 
avoid the unpredictable response to a normal 
contingency and gain the advantage of both 
parties speaking and thinking alike. 


Every branch of scientific endeavour through- 
out the world is short of trained personnel. The 
Botanic Gardens, Singapore, has not been un- 
affected. The normal compliment of qualified 
staff is four botanists, i.e. a Director, an Assistant 
Director, a Keeper of the Herbarium, and a 


Botanist, and two horticultural Curators. From 
1946 to the end of 1954, except for a short period 
in 1948-49, there was no Assistant Director. 
From 1952 to 1956, the post of botanist was 
vacant. From 1946 to 1954, the horticultural 
branch was virtually down to one officer, as one 
Curator was seconded for duty as Agricultural 
Officer. These shortages have created much 
difficulty in carrying out a full programme of 
research and have thrown additional burdens on 
the existing staff. This Department is not in the 
process of "empire-building"; its full establish- 
ment of botanical officers has remained un- 
changed for 33 years, and its compliment of 
Curators has been reduced to one since the 
Penang Botanic Gardens were handed over to 
the Department of Agriculture in 1946. 

For the first time since 1946, a full compliment 
of senior staff was at work from December, 1954, 
though one post was vacant but held temporarily 
by a re-engaged pensioner in the absence of any 
qualified younger man suitable for permanent 
appointment. For two years this position was 
maintained and even the vacant post was filled. 
Then in 1957, the Singapore Government intro- 
duced a policy of accelerated Malayanisation of 
the public service. This resulted in the loss of one 
botanical and one horticultural officer, of which 
only the latter has been replaced. Further losses 
are expected. It is not intended to discuss politics, 
but it is essential if the situation is to be under- 
stood. Briefly, it is that local men are coming 
forward in greater numbers for higher training. 
Facilities are now available at the University of 
Malaya and abroad by means of awards to qual- 
ify local men for appointment to posts hitherto 
held by officers from overseas. For the expatriate 
officer, the element of uncertainty, the offer of 
compensation for loss of career, the knowledge 
that sooner or later he will have to retire, and the 
younger he goes the better chance he will have of 
starting again in new employment are all cogent 
reasons for the loss of trained man power in 
government service in Singapore. Many ex- 
patriate officers have and are leaving before local 
men are ready to fill their posts. Some or all of 
these reasons apply to the Botanic Gardens 



though it is apparent there will be no claim for 
appointment from local persons for some years. 

Now most biological work is essentially long 
term particularly that of the botanist. Plant 
collections and written notes and papers are only 
a part of the botanical archives an institution 
possesses. Equally important are the personal 
experiences, the stored-up impressions and ideas 
of its officers. Much of this information does not 
get onto paper and published till towards the end 
of a man's career. Let me take two examples: 
The Flora of Malaya (Ridley, 1922-25) and the 
Revised Flora of Malaya (Holttum, Vol. I 1953, 
Vol. II 1955) could never have been written by 
any other authors in as complete and competent 
a form from the available collections and liter- 
ature alone simply because such works are the 
better for the authors having had first hand per- 
sonal experience. Ridley was in Malaya for 24 
years; Holttum, for 31 years. 

The point I come to then is that frequent change 
of staff where it involves a complete break of 
environment in research workers' careers is bad 
for the institution. It may be bad also for the 
individual, but that is outside the present issue. 
The changes which are now taking place in the 
Singapore Botanic Gardens through political 
circumstances are in danger of seriously putting 
the clock back. It can never, of course, go com- 
pletely back, but it does mean that over the next 
ten years or so there will be an unfortunate break 
in that intangible quality, personal experience, 
which is such an important asset to any institution. 

I have no solution to offer. Advice has been 
tendered. Recommendations have been made. 
The government has realised the situation. 
Decisions have been taken. Events are running 
their course. 

How is the loss to be made good ? The physical 
loss of bodies and the abstract loss of minds will 
no doubt eventually be recovered. For the former 
we depend on the University of Malaya and 
overseas universities where Malayans are training, 
but for the present, prospects are not good for 
only a few students are coming forward to take 
botany. For the latter, there can be no progress 
till the former is made good, and then only time 
will tell. 

This shortage of trained personnel has caused 
the introduction of a new concept of recruitment. 
Whereas it was previously accepted that persons 
arranged their own training and, except for a few 
fortunates who could obtain scholarships, paid 
for it privately, this liability now devolves to a 
considerable extent on Government and on 

public funds. Further, whereas it was usual to 
appoint a man already trained, it is now accepted 
that a promising but untrained or partly trained 
man is appointed to a post (or is anyhow bespoke 
for the post) and then trained for it. This is no 
doubt the easiest way of creating a corps of quali- 
fied man in the shortest possible time and is in 
accordance with Government's increasing con- 
cern with scientific matters. 

In the subordinate cadres, the position has been 
much easier except where specialised knowledge 
is required. Librarians appear to be hard to come 
by, and the Departmental library has been with- 
out adequate management for a long time. This 
throws out of gear all the normal services that 
one expects of a librarian such as receipts, loans, 
and exchanged; referencing and cataloguing; 
binding; pest control and prophylaxis; rack 
arrangement; and the assistance expected by 
botanical officers in locating literature. The 
solution to the shortage of librarians lies in train- 
ing, but Malaya at the present time does not offer 
any facilities and few students are going overseas 
to study for librarianships. 

One point on which this meeting might raise 
discussion is on the nature of the basic training 
required by a librarian of a botanical library. 
To run such a library, in my opinion, the librarian 
must have had some botanical training so that 
he may know the difference between, for example, 
agriculture, horticulture, and botany in their 
general meaning or between a phanerogam and 
a cryptogam in a more technical sense. With 
such knowledge he can without troubling the 
botanical staff carry out the functions of a 
librarian of a botanical library. I would therefore 
expect in the position I now find myself in the 
matter of recruiting a librarian to start with some- 
one with some knowledge of botany and to train 
him in library techniques. In this way I would 
ensure having a librarian capable of handling 
botanical literature. 

On the other hand, there is a body of opinion 
amongst librarians that a librarian should be a 
librarian and nothing else. I am assured that the 
botany around him would look after itself. But 
in a specialised library it surely calls for 
specialised knowledge to classify competently 
the constant inflow of literature, particularly if 
one considers that it is part of a librarian's duty 
to maintain a subject card index. The librarians' 
opinion to which I have just referred indicates 
a severe limitation of usefulness, but it is a posi- 
tion which now holds in Singapore and perhaps 
in general elsewhere. It is a matter on which I 
would greatly welcome comments. 



With regard to other subordinate grades we are 
in no particular difficulty for the departmental 
organisation provides the means of training 
whereby a youngster can earn and learn his way 
to positions of responsibility. In fact the Botanic 
Gardens of Singapore in some respects are rather 
like an old family concern; out of the twenty-six 
botanical and horticultural posts on the staff, no 
less than nine are filled by fathers or sons of 
present or past employees of the Department. 
I myself take pride in being in this category. 


The Botanic Gardens, Singapore, is the only 
botanical institution in the British Territories of 
Southeast Asia. It is true there are other herba- 
ria, but they tend to be specialised, with the result 
that the sphere of activity of the Department has 
been spread over the Malay Peninsula and Bor- 
neo. Recently occasional visits have been pos- 
sible to Borneo, but till 1955 these were restricted 
by shortage of staff. In Malaya, with ease of 
access, quite a different problem has arisen. For 
the past nine years there has been armed insur- 
rection in the country by a small proportion of 
the population which has taken to the forests. 
The trouble was incipient from the end of the war, 
but gained official cognisance in June, 1948, with 
the euphemistic title of "Emergency." For some 
years it reached serious proportions and even now 
is the cause of difficulties and restrictions in 
several parts of Malaya. 

This state of affairs has had a very important 
effect on collecting and botanical exploration, as 
it has been impossible to penetrate the forests. 
At times it has been forbidden, or unwise, to stop 
one's car on the main roads to have a look at the 
roadside vegetation. To wander into the forest 
would have been an invitation of trouble with 
either the police or military on patrol or from the 
insurrectionists. In either case, one would have 
been at the receiving end. Planters, miners, 
foresters, and surveyors have had to work accom- 
panied by armed escorts. 1 know only too well 
from personal experience the physical and mental 
limitation such a condition imposes. 

The consequence has been that collecting has 
been severely restricted to those areas one could 
safely visit and these on the whole are the areas 
best known botanically, i.e. the vicinity of towns 
and hill stations. The less accessible centre of 
the Malay Peninsula is where exploration is most 
wanted, for example, Gunong Kerbau, Gunong 
Tahan, Ulu Kelantan, and theTrengganu plateau. 


Armed conflict such as Malaya is experiencing 
has been or is a feature in most of the countries 
of Southeast Asia during the past decade. It is 
something with which some at this meeting may 
have had close personal experience. It is some- 
thing which all of us can deplore, though, I fear 
there is nothing this meeting can do to alleviate it. 
It has been the biggest obstacle in the past ten 
years to sociological, industrial, and research 

The situation improves in Malaya, and botan- 
ical exploration into the more remote parts of 
the country may soon be possible again, though 
the pre-war style of expedition lasting a month 
or more with men and equipment to match is 
likely to be little more than a pipe-dream except 
on rare occasions, for even some dreams come 
true! But in general we will have to work on the 
principle of light excursion lasting not more than 
a few days or on single-day collecting trips. 

An aspect of some importance is the need for 
botanists to have some familiarity with the types 
of vegetation of neighbouring countries. A 
foreign correspondent writes that he cannot plan 
a trip to Malaya as he is unable to obtain the 
necessary currency exchange. A Malayan cor- 
respondent says his proposed trip to another 
country is limited by the same curbs. I would 
like to suggest that exchange of botanists between 
institutions of neighbouring countries should be 
encouraged in much the same way as students 
and university lecturers are exchanged. The free 
provision of herbarium amenities is naturally 
simple, but the principle may well be expanded 
to the offer on a reciprocal basis of facilities for 
field work, transport, subordinate assistance, 
and lodging. 


Many scientists call at Singapore on their way 
to other countries. Few stay to work. If tropical 
botany is to be better understood by botanists 
from institutions in temperate countries, it is very 
desirable that some botanists come to work for 
short periods at tropical herbaria. They thus 
acquaint themselves with some of the problems 
of tropical botany, carry their impressions back 
to their home institutions, and interest their 
colleagues. In the last few years, we (Botanic 
Gardens and the University of Malaya) have been 
fortunate in obtaining grants to permit two young 
botanists to come to Malaya for a year each. 
It is hoped that this may be repeated regularly 
to the advantage of all concerned. There are 



certainly numberless lines of investigation ready 
for working on. The grants have come from the 
British Government Colonial Welfare and 
Development Scheme, but there are International 
Agencies which could or should furnish some 
assistance. Claims for assistance are more than 
their funds can stretch to, and botanical matters 
do not usually rank high in priority. Plants are 
fundamental to life, and their study is man's first 
science. It is our duty to raise the sociological 
status of botany in the minds of the world's 


Botanical security can be achieved only by 
ensuring permanent accessibility to collected 
material, which are the basic botanical archives. 
Essentially, the problem is one of availability and 
adequate distribution. 

The development of botany in the countries of 
Southeast Asia at the present time is still very 
much interwoven with herbaria of the temperate 
world. The connections of the Malaysian flora 
are especially strong with Kew, British Museum, 
Leiden, and the Arnold Arboretum. Many other 
links exist between tropical Asia, tropical Africa, 
and temperate countries. 

For our part, our system of distribution of 
duplicates may help to perpetuate this depend- 
ence on Europe, but it is a system which cannot 
be changed if we are to continue to make the best 
uses of our material and if we are to follow the 
principle of distribution for security sake. That 
our top duplicates go to those institutions where 
experts are working on the groups of plants con- 
cerned is a general policy. For us it usually means 
to Kew, Leiden, and the British Museum. The 
longer we send to them, the more important their 
Malayan collections become; the more important 
their Malayan collections are, the more reason we 
have for adding to them and keeping them up- 
to-date. This is a sort of cleft stick in which we find 
ourselves, albeit not an unpleasant one, nor an 
unprofitable one. This is the position which we 
at Singapore hope to maintain at least for the 
foreseeable future till there is a capacity and talent 
amongst the local population for things botanical. 
Thereafter one would wish that such distribution 
be maintained as a feature of cooperation for 
mutual benefit. The extent of distribution of 
duplicates of the higher plants after sending to 
these museums takes a wide range. All is done 
on free exchange, but it is limited to those her- 
baria of Europe, N. America, and Australia with 

Malaysian affiliations and to herbaria of the 
Indo-Pacific region, with whom we are in ex- 
change. There is, I believe, a growing opinion 
that a regional flora cannot be considered as 
peculiar to a water-tight compartment. The time 
is coming when the floras of the tropics may have 
to be taken as a whole. I am thinking, therefore, 
that our system of distribution is inadequate and 
that we may have to exchange our material for 
tropical African and tropical American material 
also, if the expanding requirements of taxonomic 
study are to be readily met. This will impose 
greater demands on storage space and the limited 
numbers of duplicates available for distribution. 
It is perhaps worth mentioning that during the 
past three years for every number collected by 
our staff and laid in our herbarium there has been 
an average of 4^ duplicates available for dis- 
tribution. We could without difficulty distribute 
double that, and we would do so were it not for 
the difficulty of handling so many duplicates in 
the field. Any increase in the number of exchang- 
ing institutions has obvious drawbacks. 

It is, however, not so much the normal run of 
current collections which give rise to the greatest 
concern, but the type specimens, the classical 
botanical archives. The Singapore Herbarium 
has about 1,000 type sheets, many are holotypes, 
others are iso-types. There may or may not be 
duplicates elsewhere; but of the older ones, it is 
likely that distribution was much more limited 
than one would now-a-days deem desirable. 
But whatever their distribution elsewhere, we have 
an obligation to ensure their availability to present 
and future botanists. Several herbaria with 
valuable collections suffered losses during the 
late war. The Singapore Herbarium was fortunate 
to have had no damage, for which the actions of 
Professors Tanakadate and Kwan Koriba, Dr. 
Holttum and Mr. Corner must be recognised. 
But preservation whether from war damage or 
from accident is a problem concerning all her- 
baria housing valuable material. For ourselves, 
we have recently had most of our type sheets 
microfilmed and propose to distribute positive 
copies to certain herbaria for their reference as 
well as for safe custody. Except for an adequate 
distribution, this is perhaps the nearest approach 
to a satisfactory solution that one could have, 
expedient though it is. 


Free exchange and free loan are, in general, 
principles to be commended. Loans from the 
Singapore Herbarium since 1946 have involved 



39,000 sheets or about 10% of the total herbar- 
ium collection. These loans, totalling nearly 
100 lots, are looked on with favour as most of 
them are for revision purposes, and the sheets 
thus come back annotated with the current 
taxonomic name. 

The increasing complexity of taxonomic revi- 
sion is reflected in the length of time these loans 
are out; for those sent out and returned, the 
average has been twenty-two months. Seven 
were out over four years; two over five years; 
and one over six years. Of the material currently 
out on loans, there are 11,000 sheets issued in 
thirty-two different consignments covering genera 
or sections or, for a few, the whole material of 
orders. Three have been out over nine and one- 
half years; two over eight years; and two over 
seven years. I cannot help but think that some- 
thing has gone wrong with these! What of the 
fervent instructions of loaning herbaria that 
the loan is only for three months? 

The accusing finger can no doubt be pointed 
at the Singapore Herbarium too. It is a common 
problem, but the point I would make is that the 
extensiveness of loans now out is apt to cause 
considerable embarrassment in handling current 
collections, as there is much which cannot be 
matched. It is not necessarily a temporary prob- 
lem, but is likely to grow with the expansion of 
botanical study. 


Most botanical institutions have attached to 
them some extent of land, garden, estate experi- 
ment station, or whatever the appropriate title 
might be, where plants may be grown under 
observation. A botanic garden usually attracts 
many visitors, though basically it is a research 
station rather than a pleasure garden. The 
Singapore Gardens were laid out originally as 
the latter on formal lines of landscaping, in all 
eighty-six acres, when in 1882 the Government 
took them over as part of a botanical institution. 
The design has remained unchanged for almost 
100 years. 

Problems inherent in the setup of the Singapore 
Gardens are these: 

(1) Space; 

(2) Pressure of the public. 

With regard to the first, the Gardens are but 
four miles from the centre of the city and are in a 
built-up residential district. Expansion is impos- 
sible. With the old Economic Garden which was 
attached to the Botanic Gardens, the area was 188 


acres which was close enough to 200 acres, 
often considered an optimum size. In 1925, the 
Economic Garden of 102 acres was excised for 
a college football field. The Frenchman's charac- 
terisation of the British "Toujours le Sport" is 
justly applied. But the result is that chiefly in 
regard to tree species, planting is greatly res- 
tricted. If the lawns of the gardens were planted 
over, more desirable introductions could be tried, 
but concession has to be paid to appearance and 
to horticultural considerations. The need for an 
arboretum is paramount at the present time. 
The acquisition of additional land has been con- 
sidered off and on for the past thirty years since 
the Economic Garden was lost and is now com- 
ing to a head again with further demands on the 

The other problem has many facets. The 
Gardens are one of the few open spaces within 
the City Area. In fine weather on week-ends or 
public holidays, many thousands of people visit 
them. In general the public's behaviour is per- 
fectly satisfactory, but in their train is the time- 
old trouble of the few anti-social individuals 
against whom restrictions have to be imposed 
which affect all visitors to the Gardens: the bicy- 
clists, who as a class, are pests; the poachers who 
try to catch fish in the Gardens lake; the pilferers 
who break cuttings ofT plants so that the plants 
have to be removed; the orchid fanciers who 
remove orchid pollenia for fertilising their own 
blooms and thus cause our plants on display to 
fade quickly. These are the troublemakers; their 
kind is common throughout the world. 

Of our own domestic horticultural problems, 
the biggest is monkeys. They are utterly des- 
tructive. All horticultural practices arc fair game 
for interference. Anything special has to be 
established under a wire cage or behind an elec- 
trified fence. Any attempt to reduce their numbers 
is a signal for outcry ; ones office is beset with holy 
matrons and true citizens defending their birth 
right, for to the public the Botanic Gardens are 
a place to come to feed the monkeys. 

The amenities which the public has in the 
Gardens are undoubtedly appreciated, and it 
seems to me that the best possible balance is 
struck between botanical/horticultural require- 
ments and the public's usage as a park. With 
more children learning biology in the schools, 
it is gratifying to have orderly parties conducted 
by teachers. This is a good augury, for there are 
many too many people who look upon plants as 
something to cut down or pull up according to 
size, and it is a function of a botanic garden, 



rather than a park, to foster a love of plants in 
the predominately destructive human mind. In 
the following section I refer to a very closely 
related subject. 


This subject is dealt with in another sympo- 
sium, but as it is a problem confronting the Botan- 
ic Gardens, Singapore, it is correctly mentioned 
here, if but briefly. 

In Singapore, the preservation of small sample 
areas of vegetation of the indigenous flora (and 
fauna) is a major problem. There are five areas 
covering about 8,000 acres constituted nature 
reserves by law under a board of trustees with 
the Director of Botanic Gardens as Chairman. 
Though this legislation was enacted as recently 
as 1951 and even more recent land utilisation 
planning has respected the areas as reserves, it is 
proving difficult to maintain them intact against 
catch-penny commercialism, demands for 
resources they contain (granite), and expanded 
development plans hitherto unforeseen. 

Two of the reserves are type localities for species 
of plants and marine animals. All are essential 
for education and research, in addition to their 
obvious attractions as places for recreation and 
open air enjoyment. Their very accessibility 
from the city adds greatly to the amenities they 
offer, but this too adds to the covetousness with 
which certain persons would seek to destroy them 
for commercial exploitation. Though there is 
a growing land hunger in Singapore, there are 
yet other similar areas of unreserved land else- 
where on the Island awaiting rational utilisation. 


None of the problems described here are in- 
superable. Some indeed are axiomatic in running 
an organisation. Some, given goodwill and 
patience, will resolve themselves. Others are due 
to force majcur. Solutions to the first two kinds 
lie in our own hands. For the third, we must 
adapt ourselves as good citizens and good 


G.A. PROWSE : Men of long proven, practical experience 
are being ousted by others with better academic records 
but less knowledge of and training in the particular sub- 
ject. This occurs where inexperienced administrators 
fail to appreciate the relative merits of experience and 
proper qualifications. Better mutual understanding 
between institutions and the authorities they serve is 

P.R. WYCHERLEY: All papers have referred to the 
shortage of trained staff. In Southeast Asia especially, 
too few students are attracted to botany; many graduates 
seek employment elsewhere, although they are sorely 
needed in botanical, agricultural, and forest research 

K. SETYODiwiRYo: Botany is a poorly paid profession 
in Indonesia, hence the few young scientists entering the 

H.M. BURKILL: Many Singapore graduates become 
teachers. We must raise the sociological status of botanists. 

E.H. WALKER: We need better advertisement of the 
importance of botanical work which should promote 
more recognition and reward for those engaged in plant 
science. These difficulties are very general. 

C.G.G.J. VAN STEENIS: The exchange of botanists be- 
tween different tropical institutes and between them and 
the temperate region herbaria, as suggested by Mr. Burkill, 
will greatly benefit the development of tropical botany. 
Tropical institutes must not be left to develop in isolation; 
personal contacts are invaluable to all. Bogor and Singa- 
pore have laid a good foundation in accommodating visit- 
ing specialists; this successful enterprise must be extended. 
Specimens are sometimes out on loan for many years, 
because the monographers are often university teachers who 

cannot devote all their time to taxonomic revision. Plan- 
ning when the specimens are required by the borrower 
will help, and herbaria should remind borrowers of the 
outstanding loan periodically after a reasonable time for 
study has been allowed. 

R. HEIM: Young botanists must be encouraged to take 
up taxonomy. Botanists must aid the establishment of 
reserves and parks to retain living specimens of the flora, 
as well as planting formal botanical gardens. The wide 
distribution of duplicate specimens cannot be emphasized 
too strongly. 

E.H. WALKER: I endorse the need for new taxonomic 
studies. Some of the problems of loaned specimens and 
exchanges of specialists could be resolved if there were 
regular conferences of the directors of botanical institutes 
to advance cooperation. 

p.s. ASHTON: The working taxonomist and collector 
must be able to visit not only herbaria but also the different 
source areas in the region, irrespective of political bounda- 
ries. A pooling of resources is needed so that botanists in 
poor countries may reap the benefit of facilities established 
by their richer neighbors. 

E.H. WALKER: There is a lack of curators in the her- 
baria I have recently visited in the Pacific area. Funds 
have been cut, and although the senior men may remain, 
whole groups of essential assistants have been unwisely 
dispensed with. Individual institutes probably cannot 
work out their own salvation. General guidance, coordi- 
nation of plans, and assistance in bringing their needs to 
the notice of the authorities arc all necessary. 

M.S. DOTY: I would remind you that we can pass reso- 
lutions for the attention of the Pacific Science Association 
or of UNESCO, whose support will add more weight to 
our conclusions. 





Department of Agriculture, Ministry of Agriculture, Bangkok, Thailand. 

Anybody may realize that Thailand, land of 
freedom and land of smiles in Southeast Asia, 
is an agricultural country. She is very rich in 
natural resources. Her soil is fertile and suitable 
for various kinds of cultivation. Rice and teak 
are famous and recognized by many customers. 
The population is too little in comparison 
with the vast area. Her economy is dependent 
on plants and plant products. 

In Thailand, as in many other "ropical countries, 
botanical knowledge which is of basic impor- 
tance is still in its initial state. Speaking in general, 
the Thai people have a great interest in plants, 
their economic value, and even the ecological 
relations. Courses in botany are offered in the 
university, but these are only for the scientific 
background of medical, pharmaceutical, biolo- 
gical, and agricultural students. The teaching 
for the advancement of Botany is still in the 
future. The Thai flora still receive scanty atten- 
tion. No botanic gardens exist in the kingdom, 
but there are a few reserved parts of the forest. 
As a matter of fact, botanical works such as a 
survey and systematic study have been made, 
but for the use of other institutions which are 
considered the main arteries of the country's 
economy, i.e., Royal Forest Department and 
Department of Agriculture. 

In 1906, the Royal Forest Department started 
the systematic study of plants, but these were 
mainly timbers of economic importance and 
some other forest products. A herbarium was 
also built up. In 1920, the Botanical Section of 
the then Ministry of Commerce was formed 
with the object, in the first place, of making a 
survey throughout the kingdom of plants yielding 
or likely to yield, economic products. This sur- 
vey was to include an enquiry into the properties, 

quantities, and accessibility of such plants. 
Again the herbarium was built up. This Section 
is now under the Division of Plant Science, Depart- 
ment of Agriculture. A great deal of work has 
been done on the flowering plants of Thailand, 
but it is still incomplete. Studies of ferns and 
mosses have been made to a lesser extent, but 
the lower plants, fungi, and algae are almost 

One might question why botanical studies and 
work have not developed further in Thailand. 
Shortages of trained scientific personnel and 
scientific literature as well as inadequate funds are 
all responsible. 

The most important of these is the shortage of 
personnel. In the two mentioned botanical 
sections, there are only a few members of the 
official staff, and work progressed very slowly. 
Collections which are necessary and important 
are made on a small scale. There are too few 
duplicates to offer to other institutes. The main 
factor is the difficulty of finding persons interested 
in making a career of this purely scientific work. 
There is a demand for persons in various fields 
of applied science such as medicine, engineering, 
architecture, agriculture, etc. Pure science has 
thus far not attracted sufficient students and 
knowledge of the country's vegetation is still very 
scant. The shortage of trained personnel creates 
other problems in developing this botanical 
knowledge such as limited advanced study and 
little coordination with other institutions. 

At present official persons in various fields 
have begun to realize the importance of botanical 
knowledge. We hope that in the future this kind 
of purely scientific work will have the full support 
of the government, as a part of the country's 
economic development. 


C.G.O.J. VAN STEENIS: The lack of a botanic garden on 
a scientific basis in Thailand is amazing in view of the 
agricultural nature of the economy. Indonesia and Malaya 
are examples of how botanic gardens, with their herbaria 
and libraries, have aided agricultural development by 
knowledge of the flora. The establishment of a botanical 
garden requires careful long-range planning, for slow 
but sure growth is needed. This is a government responsi- 
bility. I propose that this section endorse any resolution 


supporting the establishment of a botanic garden by the 
Thai Government. (There was general agreement that a 
botanic garden is an urgent necessity for any tropical 
country, including Thailand, which lacks such an institute 
for the botanical study and development of its agricultural 
and forest resources.) 

M.L. STEINER: Our attempts to rehabilitate the gardens 
in Manila have failed. The trees and shrubs we planted 
are destroyed. The promises of help from the authorities 


have come to nothing, how can we press the point with tion is needed. The government may stand in need of 
them? education in this matter. 

E ' H - WALKER: Vigorous action is needed on all fronts. 

interest larger organizations in obtaining effective support 
C.G.G.J. VAN STFTNIS: Government, not private, ac- from the authorities. 





Flora Malesiana Foundation* Oegstgeest* Netherlands. 

Not long ago I was struck by a plea to mono- 
graphers by Brother Alain (Havana) who com- 
plained of the fact that monographers frequently 
study less material than desirable and often 
confine their work to the revision of the material 
of some large, European herbaria (Taxon 6, 
1957: 46-47). He pointed to the deficiency in 
this respect of Radlkofcr's monograph of the 
Sapindaceac in the Pflanzenreich. 

It is true that in many monographs of this 
series only a very limited amount of material 
was borrowed from institutes outside Berlin- 
Dahlem, and that for essential material only. 

Two decades ago when working at Bogor, 
Java, I had a similar complaint about Malaysian 
material, and I wrote to Berlin to ask if we could 
receive timely notice as to which families or 
genera were under revision in order to loan our 
material for examination by the monographer. 
The answer was unsatisfactory, as it was argued 
that the monographers could generally not 
master large loans from all herbaria abroad and 
did not solicit to ticket all specimens, as their 
facilities and specially their time were restricted. 

Nearly all of the monographs therefore suffer, 
in varying degree, from incompleteness. And 
the same holds for various other revisions general- 
ly. The reference to Radlkofer's work is not 
a particularly good example as it was essentially 
finished before 1910 and remained, as far as I 
know, in Radlkofer's desk for decades. Radl- 
kofer did not add much to it in later years, and 
the final posthumous editing was performed by 
the redaction of the Pflanzenreich. 

This situation looks lamentable and dangerous 
to the progress of international taxonomical work. 
It is partly due to the fact that tropical collections 
are accumulating so fast that many large herbaria 
are overcrowded. Furthermore, the number of 
herbaria is still increasing. 

On the other hand, monographers are still 
scarce or monographic work is not sufficiently 
envisaged to be the major source of creative work 
and the available time of each monographer, 
in contrast with the dynamism of collections, re- 
mains static, i.e. twenty-four hours in a day and 

A long experience with the Flora Malesiana 
revisions, which cover only a part of the tropics, 
has taught that completeness in ticketing all sheets 
which is generally a multiple of all collecting 
numbers of all herbaria in the world containing 
Malaysian plants is an impossibility. And from 
this experience I infer that such a procedure 
for a monograph of even only a medium-sized 
family would lead to insurmountable personal, 
technical, and financial difficulties; it requires 
an immense amount of time from the side of 
the monographer to cope with the routine work 

To give a slight idea of what colossal numbers 
of sheets must be revised for a monograph, in a 
study of Cyperus only for Malaysia and from a 
limited number of herbaria, about 17,000 sheets 
passed through the hands of Mr. Kern for 
comparison or identification and ticketing! For 
Fimbristylis, the number of sheets was about 

The historical development of plant taxonomy 
and distribution of collections has led to the 
present situation of rather few large world her- 
baria in which collections are presented and 
worked on from all parts of the world, and rather 
few very large herbaria which are more or less 
concentrated on certain major regions of the 
World (Africa, New World, Old World, Austra- 
lia, etc.). Whether we like it or not, this situa- 
tion has to be accepted. 

Most of these institutes have less scientific 
staff members than would be desirable; some are 
even ridiculously understaffed. Generally staff 
members must, by necessity, perform official 
and routine work and are left little time for mono- 
graphic work. 

Combining this need of time on the side of the 
monographers with the just mentioned immense 
increase in sheets, there is an absolute necessity 
for them to use their time as efficiently as possible, 
resulting in limiting their work primarily to 
essential material and to that which is immediate- 
ly available, rather than to strive towards com- 
pleteness in ticketing all specimens of their group 
they can lay hands on. After all, a revision or 
monograph derives its value from its usefulness, 



from keys that work, from the degree of accuracy 
by which genera and species are delimited and 
described, and not from immense lists and 
enumerations of specimens from this or that 

I have touched on this subject in my prepara- 
tion of the Flora Malesiana (ser. I, vol. 1, p. 
Ixv) in a chapter on the policy of distributing 
duplicates as advocated and applied by the late 
Dr. Merrill in the Philippines who was in a simi- 
lar position as we were at Bogor and Brother 
Alain in Cuba. Dr. Merrill has recommended 
that the tropical collectors and institutes take the 
initiative and distribute duplicates to the large 
world herbaria, specially to those from which it 
can be expected that they will name them immedi- 
ately or in future, and to those where already 
large, authentic collections are represented from 
the region where the new collections are made. 
Of course the tropical institutions cannot expect 
to come on terms of an exactly balanced ex- 
change, as the world herbaria have generally 
little to offer in this respect. Dr. Merrill did this 
always on what he called a "free exchange basis," 
that is: both parties give what they are able to 
produce. As far as material is concerned, the 
tropical institutions will therefore be largely the 
donating party. 

But their effort is materially rewarded in several 
respects: firstly, the collections, provided they 
have been preliminarily identified to families and 
preferably to genera, come into "scientific cir- 
culation," they arc "available" and will come more 
easily under the eyes of monographers, interested 
staff members, or visitors of the large institutes. 
Secondly, wide distribution of duplicates safe- 
guards collections against calamities. Thirdly, 
the reward to the donating party will exist in the 
ultimate naming of its collections by specialists. 
Fourthly, its initiative and diligence provides the 
donating institution with favourable contacts 

abroad and with specialists; it will stir up interest 
in tropical plants. In passing I may give the 
practical hint of printing at the bottom of each 
duplicate label a request that any new identifi- 
cation of the sheet be kindly brought to the 
notice of the donating institution. 

The history of the exploration of the Philippine 
flora, through the impetus of Dr. Merrill, proves 
the wisdom of the policy just mentioned. Even 
though the calamity of war came over Manila 
and destroyed what had been built during two 
decades of extremely hard, intelligent, and dili- 
gent work, the liberal way in which duplicates 
had been distributed still guarantees the possibil- 
ity of writing a complete Flora of the Philippines. 

The rich collections assembled in tropical 
centres must be brought to the notice of the work- 
ers and visitors of the big world herbaria. This 
is the duty of tropical assembling places, and 
besides a duty it is a great privilege, because trop- 
ical botany will, for at least half a century to come, 
not only be mostly dependent on, but will also 
benefit from contributions of the great scientific 
centres in the temperate regions of the globe. 

I believe that Brother Alain has touched on a 
subject worthy to explain during this Congress 
which serves to cultivate goodwill, understanding, 
tolerance, and cooperation. On one hand I be- 
lieve he underestimates the labour involved in 
monographical work; on the other hand, he un- 
derestimates the extreme value of tropical herb- 
aria taking the initiative in their own hands. Trop- 
ical herbaria should recognize the extreme value 
of a very liberal policy of distributing duplicate 
specimens of good material of native plants to the 
world herbaria. Specialists, on the other hand, 
are invited to give timely notice, preferably in 
Taxon, of the subject of their intended monograph 
or revision. In this way a satisfactory interaction 
and cooperation can be realized. 


E.H. WALKER: The collection at Manila was destroyed 
by fire during the war, fortunately many duplicates are in 
the Smithsonian Institute. This illustrates the need for 

wide distribution of duplicates. Breakdowns in the loan 
of this material call for more cooperation between directors. 





Fish Culture Research Station, Batu Berendam, Malacca, Federation of Malaya. 

While a botanical department of a freshwater 
fish culture research station is not strictly speaking 
the same thing as a botanical institution, many 
of the problems are similar, while others will be 
peculiar to the station itself. For that reason it 
seems worthwhile bringing these problems before 
a meeting of this kind, with the hopes that the 
ensuing discussion may result in their solution, 
or at least suggestions of their solution. 

Taxonomy. Limitations of staff make it neces- 
sary that a botanist working in such a station as 
ours undertakes taxonomic studies on the various 
plant groups, although unlike other botanical 
institutions, he is at least limited to those plants 
in an aquatic environment. The botanist here, 
however, cannot afford to confine himself to one 
particular group and must spread his efforts, 
although inevitably he finds himself taking a 
greater interest in one of the groups. Because 
of this he finds himself dependent on the goodwill 
and cooperation of others elsewhere, and various 
problems necessarily arise. 


In common with other institutions, a newly- 
opened iish culture research station always finds 
itself short of the relevant reference works. This 
is as true in the botanical department as elsewhere. 
Many essential books are scarce or out of print, 
while others are so expensive as to be beyond 
the department budget. It is vitally necessary, 
particularly when describing a new species, that 
the botanist should have all the available reference 
works at hand. The lack of this is seen in the 
appalling number of synonyms for many organ- 
isms. Photo copying seems to be the answer to 
this lack of relevant literature, and it can take 
several forms. Photostats are large and easily 
read, but they are comparatively expensive and 
are better bound for storing. Microfilms are 
cheaper but necessitate a special reader and are 
troublesome to store, and it is not always easy to 
refer to a page and illustrations at the same time. 
Microcards, where they have been printed, are 
much better, and are ideally used with a special 
reader but are quite easy to read with a dissecting 
microscope. Furthermore they are black print 
on white background, and are easy to store in 


filing cabinets, being the same size as a filing card. 
In cases where microcards have not been made, 
microtape can be made from an existing micro- 
film and gummed on to a filing card to make what 
is in effect a microcard. The extra cost is very 
little. All these methods may be beyond the 
budgetary limit when numerous large works have 
to be copied. 


Lack of literature at hand means that either 
we must borrow, or else send material elsewhere 
for identification. With the macrophytes, par- 
ticularly the flowering plants, good herbarium 
specimens should present little difficulty. With 
the microscopic plants, however, especially the 
algae, numerous special problems arise. Drawings 
of the organism, in as many aspects as possible, 
or good photographs are often valuable, but 
there is always the danger that the artist's inter- 
pretation is not strictly accurate, and a photo- 
graph always lacks depth of focus. Nevertheless, 
in the absence of the actual organisms themselves, 
these can be a very good second best. 

The despatch of actual material is a little more 
difficult and entails special methods. The usual 
method of preserving algae is in 5% formalin, 
and for things like desmids or diatoms, where 
the structure of the cell wall is important, or for 
the Chlorococcales, it seems perfectly adequate. 
For cell contents such as in Spirogyra, it often 
causes distortion, while the more delicate flagel- 
lates may become almost unrecognisable, and 
colonial forms may break up. In such cases 
Picric acid, or one of the compound preservatives 
might be better and it would be necessary to 
experiment. The flagellates in particular are a 
difficult problem as they so easily become un- 

Permanent slides suggest themselves as an 
answer, and they are particularly important for 
the preservation of holotypes. The mounting of 
the frustules of diatoms is too well known to cause 
much concern. With other algal groups it is not 
so easy. The serial dehydration necessary for 
many mountant fluids often leads to the distortion 
or complete loss of the organism concerned. 



A direct mountant is indicated, and in this respect 
I have found Polyvinyl alcohol very useful. It 
both fixes and mounts the living organism and can 
be diluted with water to the required strength to 
prevent plasmolysis. It will set hard, although in 
the wet Malayan climate I have found it better to 
encourage its drying-out in a desiccator. Its 
main disadvantage is that objects become so 
transparent in it, and it may be necessary to add 
a dye. Glycerine mounts, while easily prepared 
and useful, are too fragile to send by post. More 
experiments are obviously needed to find the best 
permanent mountants for the algae. 

The ideal way would be to send living material, 
for then it could be examined in the living state 
and even cultured. Naturally this presupposes 
despatch by air, and if it is possible to send the 
material direct from airport to airport without 
intervening postal delays, so much the better. 
The problem is to keep the organisms alive and 
in reasonable condition throughout the journey. 
Drastic variations in temperature are nearly 
always harmful to algae, and these are just what 
they seem to be subjected to when carried from 
country to country. To overcome that, it would 
be necessary to insulate the container as much as 
possible, or even enclose it in a thermos flask, 
with consequent increase in air-charges. As 
containers for the water samples, 1 have found 
polythene bags very useful, as there is little danger 
of them breaking, and several of them could be 
packed inside a single larger container. Another 
problem is bacterial action, which can become 
quite serious in the time taken for the sample to 

reach its destination. Field collections of water 
samples inevitably contain some bacteria, and 
these are certain to multiply. Perhaps the careful 
use of one of the antibiotics may overcome this 
difficulty. A further serious difficulty is oxygen 
deficiency in the water. It must be remembered 
that the algal specimens will be travelling in the 
dark, and therefore any oxygen used up in their 
respiration cannot be replaced by that due to 
photosynthesis. Furthermore, in natural field 
collections there are bound to be a number of 
small animals which will further deplete the 
oxygen supply. The tubes or polythene bags 
should not be more than half full, and any dense 
suspensions should be diluted with well oxy- 
genated water. Living samples have been suc- 
cessfully sent over long distances, but they have 
usually been pure cultures with very few organisms 
in the container. Thus if a pure culture, partic- 
ularly bacteria- free, could be maintained, it 
would be no difficulty to put a minute drop in 
\ tube of sterile culture solution. The estab- 
lishment of such cultures, while essentially part 
of the work of a fresh-water fish culture research 
station, is outside the scope of this paper. 

The foregoing should give some idea of the 
problems which occur in such a specialized 
botanical department as ours at Malacca, and 
I have no doubt there are many other such de- 
partments with similar problems. I have indi- 
cated some of the possible ways of solving these 
problems, but I am hoping to gain much informa- 
tion and advice from the discussion which will 


E.H. WALKER: Some of the problems are bibliograph- 
ical, and I hope, as I have mentioned from time to time, 
you will support the meeting on this subject. 

M.S. DOTY: Polythene bags are permeable to air, 
although not to water. The amount of water may be 
profitably reduced until it forms only a thin film which can 
absorb the maximum oxygen. 

G.A. PROWSE: Desmids and flagellates are the most 

M.S. DOTY: Some fixatives are of value. 

E.H. WALKER: Cooperation between institutes and 
airlines are needed to improve transit facilities. 

H.M. BURKILL: The use of the diplomatic bag has been 
advantageous in difficult cases. 

E.H. WALKER : We need scientific attaches. 

J.H. HORLIMANN: They could help in the numerous 
quarantine difficulties. 

p. WEATHERWAX: Soil samples are often unacceptable 
to the health authorities, but they can be sent through with 
cooperation. What are conditions like in the freight holds 
of aircrafts? 

M.S. DOTY: Specimens must go in the pressurized part 
of the plane; the freight holds are freezing. Packages 
should be clearly labelled to this effect, and, if possible, 
be put in the hands of the second officer of the aircraft, 
who can also reduce delays or unfavorable storage on the 
ground at intermediate stops. 





Museum National (VHisloire Naturellc, Paris, France. 

This note aims essentially at suggesting a 
few measures which would enable the conditions 
of botanical research within our territory of the 
Southwest Pacific Islands to be improved. 


In this field, efforts should be directed, above 
all, towards enabling the investigator who is not 
always an expert to have the means of making 
a rapid and exact determination of the specimens 
he has gathered or, inversely, towards providing 
him with a precise representation of the morphol- 
ogy of the species he wishes to obtain. 

In this connection, it should first of all be made 
clear that no adequate reference herbarium 
exists on the spot at present. The material which 
can be utilized is limited, in this case, to fragmen- 
tary collections provided exclusively by metro- 
politan and foreign botanists. The investigators 
thus find that they arc almost always obliged to 
have the necessary determinations made, or 
confirmed, by the competent services of the 
Natural History Museum in Paris, this leading to 
a considerable loss of time. As a palliative to 
this difficulty, the following measures would 
appear to be highly recommendable: 

a) the drawing up of several basic herbaria, 
as complete as possible, centralised in Nou- 
men at the seat of the scientific and adminis- 
trative bodies who are the most directly 
interested: Institut Frangais d'Oceanie, 
Commission du Pacifique Sud, Service des 
Eaux et Forets, Service de T Agriculture, 
Mus<e no-caledonien. Botanists who are 
collectors "de passage" should always under- 
take to enrich these herbaria by the deposit 
of duplicates; and where only one specimen 
of a new species is gathered, this should be 
provisionally represented by a full-sized 
photographic reproduction; 

b) the opening, for each species, of an identical 
file, the number of these corresponding to 
the basic herbarium and which will contain, 
in addition to the classical elements, a very 
detailed morphological description, a syn- 

t Presented by J.H. Hiirlimann. 

onymy, vernacular names, general, chorolo- 
gy, list of the known neo-caledonian locali- 
ties, ecology, etc. A photographic documen- 
tation carried out in the natural state and 
referring not only to the leguminous aspect 
as a whole (port), but also to details: leaves, 
flowers, fruits. 

Another problem, of primary importance, is 
that of a practical bibliography. The only flora 
published up to the present- that of A. Guillau- 
min (1948) was intended by the author as a 
preliminary work only, destined to meet imme- 
diate needs. Consisting of synoptic tables only 
and totally deprived of illustrations, it is accessi- 
ble only to investigators who are already well- 
informed. In addition, the rigid framework of its 
presentation does not lend itself easily to am- 
plifications, a disadvantage which is particularly 
inconvenient when one realises that there still 
remains a great deal to be done before the floristic 
inventory of New Caledonia is complete. In our 
opinion, the ideal formula to adopt is to be 
found in the drawing up of a flora carried out on 
the principles which inspired O. Degener for the 
publication of his New Illustrated Flora of the 
Hawaiian Islands: a perforated sheet movable 
for the description of each species, the recto 
being reserved for the text (diagnosis, ecology, 
chorology, etc.) and the verso for several sketches 
of their features. It is clear that such an arrange- 
ment would allow for the easy insertion of addi- 
tional sheets concerning new species in the same 
way as it would allow for the replacement of 
obsolete keys of determination by up-to-date 


Cartography. For the moment, the topograph- 
ical Services of New Caledonia put two types of 
terrestrial maps at the disposal of the public: 
1:40,000 maps and 1:10,000 maps. Utilized 
specially for the delimination of mining and 
agricultural concessions, the 1 :40,000 maps cover 
only a fraction of the area of the Grande Terre. 
Further, their planimetric representation is 



simplified in the extreme. Indeed, we find only 
the contours of the coast and the waterways and 
roads, with a few summary particulars with regard 
to swamps, the European settlements, the large 
forest areas, etc. As regards the relief, dotted 
lines indicate the position of the principal ridges 
and the peaks mentioned are mostly defined, not 
by their altitude, but by their order number with- 
in the triangulation framework. Although the 
1:10,000 maps in curves are of course more 
exact, they exist only in scattered sheets and 
cannot therefore serve for purposes of general 

However, all these gaps are shortly to be satis- 
factorily solved, thanks to the publication of a 
new map under the auspices of the Institute 
Geographique National. Drawn up from aerial 
surveys, this map, the final scale of which is not 
yet fixed (1 :50,000 or 1 : 1,000,000), will cover not 
only the Grande Terre as a whole but also the 
islands geographically dependant on it: Pine 
island, the Loyalty group, etc. 

We would also mention the excellent geological 
map 1:100,000 in 10 sheets (Grande Terre and 
Pine Island) drawn up following on the work of 
the mission led by P. Routhier from 1946 to 
1949. Already partially on sale, within the 
programme of publications of the Office de la 
Recherche Scientifique et Technique Outre-Mer, 
it represents, a considerable advance compared 
with our previous knowledge in the subject and 
will be of inestimable service to the ecologist. 

Ecology. While the equipment of the pedolog- 
ical section of the French Institute of the Pacific 
Islands now enables very complete analyses of 
soils to be carried out, nothing positive has as 
yet been accomplished in the field of micro-cli- 
matological researches. The local National 
Meteorological Service whose observation sta- 
tions are, incidentally, almost all placed in the low- 
lying regions near the shores directs its activities 
mainly towards objectives that are primarily 
practical: definitions of the characteristics of the 
general climate, weather forecasts with a view to 
the improvement of the conditions of air and 
maritime traffic, etc. 

The accelerated developments of the ecological 
discipline requiring data which are more and more 
exact, from now on the serious study of the 
dynamics of the leguminous groups of New 
Caledonia, cannot be envisaged without the 
support of more solid arguments than the spora- 
dic measurements carried out up to now by na- 
turalists in the course of their movements. In 
the United States, for instance, the use of auto- 

matic and portable micro-climatological stations 
placed within the various strata of given groups 
has already given very encouraging results. 
Noted every week, the tapes of the registering 
instruments thus constitute homogeneous series 
of continued observations. This technique, it 
seems to us, should find a perfect application 
within the limits of our Southwest Pacific terri- 
tory, more particularly within the upper zones 
of the mountainous regions where it is not always 
easy to remain for sufficiently long periods. 

Transport. At the present time the roads ac- 
cessible to motor traffic are limited to two arterial 
roads running along the East and West Coasts 
and communicating with one another by three 
or four transversal roads. In very many cases this 
arrangement hinders, in particular, the rapid 
approach to areas of work situated in the moun- 
tain groups. Very often, long and fatiguing 
distances over extremely varied ground have to 
be covered on foot in order to reach the goal 
which has been set. During the ascension, the 
investigator, obliged to mobilise all his physical 
forces in overcoming natural obstacles, can devote 
only a small part of his intellectual faculties to 
detailed scientific observation. Heavy rains or 
intense sunshine often add to this depressing 

In the absence of suitable landing ground, 
the light airplane would not be suitable to radical- 
ly overcome these difficulties, but the helicopter, 
to which the problem of a large landing ground 
does not apply, certainly represents the solution 
for the future. Incidentally, one of these helicop- 
ters used experimentally by a local industrial com- 
pany for mining prospection has, up to the 
present, given entire satisfaction. Thus put down 
in the area of work without any previous fatigue, 
the investigator finds himself able to give of his 
very best during the time he remains on the spot 
and, afterwards, also during the descent towards 
the lower lying regions. 


The transport difficulties of which we have 
just spoken have very considerably retarded the 
inventory of the neo-Caledonian vegetation. 
Indeed, great progress still remains to be made in 
the knowledge of nature and the distribution of 
the leguminous groups. In order to attain this 
end, it will be necessary to call in the assistance of 
aerial photography on a large scale and in colour. 
Personally, in the course of flights at low altitude, 



we noted the clearness with which certain masses 
physiognomically very individualised, such as 
the Mangrove, the herbaceous belts of the sandy 
beaches, and the photophileous forest of Melaleuca 
Leucadendron, were to be distinguished, thanks 
to their morphology and their colouration. Of 
course, a general map, made up from an assem- 

bly of aerial cliches, does not in any case do away 
with the classic analytical surveys made on the 
spot; but, judiciously interpreted, it possesses the 
enormous advantage of enabling us to give the 
precise localisation of the large laguminous for- 
mations and, as a consequence, to rationally 
coordinate the researches on the ground. 


M.S. DOTY: I fear helicopters may prove out of the 
question; it costs US$400 to fly one for an hour. 

J.H. HURLIMANN: I admit we may not be able to realize 
all our hopes, but the possibility is noted. 






Division of Botany, Department of Forests, LAE. 

The collection of botanical specimens in 
tropical countries has always presented a number 
of problems not faced by collectors in temperate 
latitudes. The normally high humidity combined 
with temperatures usually exceeding a minimum 
of 75 degrees Fahrenheit bring with them the 
problem of rapid fungal and bacterial growth. 
A large number of tropical plants, particularly 
those of the rain forests, develop abscission tissue 
with extreme rapidity in the leaf petioles and 
petiolules. An inspection in any herbaria of the 
specimens of tropical plants will reveal how 
widespread this is. 

Various botanists in the tropical woHd have 
met these problems in a variety of ways. Eleven 
years experience in Papua and New Guinea has 
produced the techniques and procedures which 
will be described below, but before doing so it is 
necessary to give some background regarding the 
nature of the country, its people, and tran- 

In New Guinea, botanical collecting has been 
and still largely is associated with geographical 
exploration. As yet little has been done toward 
the exhaustive compilation of the flora of a 
limited area. The botanist and his equipment 
must therefore be mobile. Road transport is 
available to a limited but rapidly increasing 
extent. Air transport particularly by light aircraft 
carrying loads up to 1,200 pounds weight is 
widely available. Water transport descending 
in size from coastal vessels of about 300 tons to 
canoes is generally available in coastal areas. 
Canoes and small launches may be used to some 
extent on most lowland rivers. A few areas may 
be reached by float plane, landing on lakes. 

Using the most appropriate transport, the 
botanist gets himself into the general area he 
plans to visit. Further progress is on foot with 
human porterage as transport for the collecting 
equipment. Fortunately, in most areas the local 
inhabitants are prepared to assist parties by 
carrying equipment and supplies. The rate of 
pay varies rather widely throughout New Guinea. 
The maximum load is about 40 pounds per adult, 
or where a load is carried jointly by two men, 

t Presented by H.M. Burkill. 

a total of 80 pounds weight. Equipment must 
therefore be of a size and weight which allows it 
to be made up into suitable sized bundles for 

The actual equipment used differs somewhat 
from that of a botanist in temperate countries. 
Most of the flora is woody so that an axe is 
indispensable if justice is to be done with this 
great group of plants. One is led to suspect that 
at least as far as New Guinea is concerned the 
difficulty of falling large trees has sometimes 
turned the attention of botanists away from the 
tree storey. Lane-Poole, when collecting in New 
Guinea in the 1920's and being almost exclusively 
concerned with the forest flora, used a rifle 
successfully to bring down branches of forest 
trees. A shot gun can also be used to good 
advantage at times. Secateurs are extremely 
useful for cutting the actual specimens, while 
heavy gloves or a machete facilitate the collection 
of thorny palms and trees. 

As the specimen is finally prepared, usually 
represents but a fragment of the original plant, 
adequate descriptive field notes are essential. 
The value of the collections is greatly enhanced 
and frequently identification facilitated if wood 
samples are collected. General practice in New 
Guinea is to obtain a wood sample of all plants 
having a woody stem. If this exceeds 6 inches in 
diameter, the sample is usually trimmed to 15 
inches long with a cross section of 4 inches by 
4 inches. Bark is retained wherever possible. 
Where weight is a serious consideration, the wood 
sample may be reduced in size. 

Turning to the specimens of foliage, every effort 
has to be made to collect flowering and fruiting 
material of each species. Most tropical herbaria 
have a considerable quantity of sterile or inade- 
quately fertile material which can be only partially 
identified and must await the recollection of more 
adequate material. This does not mean that 
under certain circumstances such as a collector 
in a "new" country or where a plant appears to 
have some special interest, specimens lacking 
flowers or fruit should not be collected. I can 
repeat, though, that a wood sample in such cases 



can be of immense value in identifying the 

The technique adopted for preparing the 
botanical specimens in Papua and New Guinea 
is based on the formalin technique of Schultes, 
used successfully by him in South America. 
However certain modifications have been made. 
At the base camp are required newsprint, folded, 
but preferably clipped to 18 inches by 12 inches; 
frames, 18 inches by 12 inches constructed as a 
lattice from wood 1 inch by inch thick; a 
rectangular tank, the inside dimensions of which 
are 18-^ by 12} inches by 12 inches deep, 
fitted with a lid; strong string; sisalkraft; (a water- 
proof bituminous paper); and concentrated 

Actual collecting requires newspapers, frames 
(or wire press), strong cord (leather straps 
deteriorate too quickly), and labels. The speci- 
mens, suitably pruned to display the essential 
parts to best advantage, folded in the case of large 
or compound leaves, are numbered by attaching 
a number bearing manilla tag, the number on 
the tag corresponding of course to the relevant 
number in the field note book. Soft pencil or 
crayon is most suitable for numbering on the tag. 
Do not use ordinary ink. A pad of folded news- 
paper is opened so that all but the lowest sheet is 
folded on the left. The lowest sheet is reversed 
so that the fold is on the right. The numbered 
specimens of each collection are now placed 
singly between the sheets. Wherever possible, 
ten or more replicates of each collection should 
be obtained. With this number it is convenient 
to confine a single packet of paper to each col- 
lection. The reversed bottom sheet is finally 
folded around the packet thereby providing 
closure at both edges. The packets are then 
placed between a pair of wooden frames and 
pressure exerted by means of several turns of 
strong string at each end. A bundle not exceeding 
twelve inches in thickness is desirable. These 
bundles are then transported to the base camp. 

The next step consists of preserving the speci- 
mens from fungal and bacterial action by immers- 
ing them in formaldehyde solution of 4% con- 
centration by weight, specific gravity about 1.014 
at 75 degrees fahrenheit. The formalin can be 
prepared by diluting a concentrated formaldehyde 
solution. As normally received in New Guinea, 
this rarely exceeds 35% formaldehyde by weight 
and should be broken down in the proportion of 
one part of concentrate to eight parts of water. 
An alternative procedure has recently been 
adopted with success using the solubility of 

paraformaldehyde in hot water made alkaline 
with hexamine. This produces a formaldehyde 
solution, the specific gravity of which is adjusted 
to 1.014 after cooling. Solution can readily be 
effected by adding about one ounce of hexamine 
to three gallons of water which is heated to the 
boiling point. After removing from the fire, 
slowly stir in three pounds of powdered para- 
formaldehyde. Due to the formaldehyde vapour 
produced, this can be most unpleasant. An 
alternative which works equally well is to place 
the paraformaldehyde powder at the rate of one 
pound per gallon in a one gallon, wide mounthed, 
plastic, screw top bottle. The hot water, which 
contains hexamine at the rate of one ounce to 
three gallons of water, is then added to each 
bottle, the lid screwed, on and the bottles shaken 
intermittently until solution is complete. After 
cooling, the specific gravity is adjusted as before. 
The use of paraformaldehyde has obvious 
advantages over concentrated formaline where 
weight is a factor to be considered. 

Having prepared the formalin solution, the 
rectangular tank, the inside of which has been 
coated with bitumastic paint, is half filled with 
4% formaldehyde. The prepared bundles of 
specimens referred to above are immersed, a 
stone may be used to stop the bundle floating, 
the lid put on and the whole lot left for 18-24 
hours. The following day the bundle of speci- 
mens is removed; the free liquid is drained off; 
and it is securely wrapped in waterproof sisalkraft 
paper. In this condition, specimens have been 
kept without deterioration for three months. 

The bundles of formalin preserved specimens 
are sent by whatever transport is available to the 
divisional headquarters at Lae. Here the bundles 
are opened, the specimens placed between drying 
pads which are interspersed with corrugates for 
ventilation, and dried in cabinets heated with 150 
watt globes. Drying is complete in 24-36 hours. 
Where succulents or palms having a waxy epider- 
mis are being collected, the addition of a liquid 
detergent to the formalin at the rate of about one 
tablespoonful to each gallon is advantageous in 
accelerating the penetration of the leaf tissues by 
the formalin. Probably no harm would accrue 
from the general use of a little detergent with the 

The main attraction of this method as used 
in New Guinea is that the collector does not have 
to spend time while in the field in tending drying 
ovens or racks. For the successful use of the 
method, though, access to some central drying 
facility is essential. 




C.G.G.J. VAN STEENIS: JFor herbarium work, it is essen- 
tial that the morphological characters of the specimen 
should be well preserved. Dehiscence of leaves, leaflets, and 
floral members must be prevented by rapid killing. Alcohol 
and/or formalin are excellent, but whatever facilities are 
available such as heat may be used if circumstances dictate. 
I agree with Drs. Prowse and Walker, field characters must 
be recorded at collection. Otherwise such valuable infor- 
mation as scent, color, and glossiness will be denied to the 

herbarium botanists who compile the floras for practical 
use. It is essential that every duplicate specimen should 
have a full copy of the original field label. 

E.H. WALKER: The original field labels on the Manila 
specimens were destroyed in the fire; the duplicates lack 
this information, which is now permanently lost. Hence 
the need, despite the labor involved, of providing every 
duplicate specimen with a copy of the field notes. 



Symposium: Vegetation Types of the Pacific. 


Institut Botanique, Universite de Montreal, Montreal, Canada. 

Many methods are used for describing vege- 
tation, although much of the available literature 
is appallingly unmethodical in the sense that the 
standards are poorly defined. No valid criticism 
of method or approach, on the other hand, can 
be made without an honest effort to evaluate 
purpose and scope which are an ultimate measure 
of adequacy. 

This symposium on vegetation types of the 
Pacific does not hope to provide a complete 
inventory of all synecological work, even less an 
accurate description of all known plant com- 
munities. Research on vegetation has been very 
widely scattered and many bioclimatically im- 
portant areas have not been studied at all. Nor 
was it possible to request a uniform approach 
of all the participants in this joint effort. It has 
seemed more useful to pick contributors who had 
a good deal of field knowledge and who had 
already made a contribution to the description 
and interpretation of Pacific landscapes and to 
give them a free rein in their own course. The 
resulting papers arc sure to be somewhat hetero- 
geneous, and their divergence in method will no 
doubt be pointed out by Frank E. Egler. 

It seems to me, however, that some reflections 
on structure variation are an appropriate preface 
to this presentation and that examples from 
various parts of the Pacific can be usefully quoted 
in view of subsequent correlation with the geolog- 
ical, historical, climatic, dynamic and other 
interpretations which are forthcoming. At pre- 
vious congresses (New Zealand, 1949, Philippines, 
1953), I have made similar contributions which by 
now have been amply discussed with many 
colleagues. I feel that a further step can be 
undertaken at this time, and I will therefore 
offer some elaborations upon the system which 
I have previously proposed (1951, 1952, 1953, 
1957a, 1957b). 


Spatial distribution of the biomass is a function 
of life-form, size and coverage, which in turn 
may vary seasonally (relative deciduousness) and 

t Presented by F.E. Egler. 

which may be caused by diverse assemblages of 
leaf-type and texture classes. Table 1 repeats 
my earlier attempts (1951, 1952, 1957a)to provide 
a simple, practical scale for grading actual stands 
of vegetation or for abstracting therefrom a 
number of well-individualized types. 

1 have previously discussed in some detail the 
choice of the alternates presented under each of 
the six criteria that appear in Table 1. But I now 
find that systematic application by myself and by 
others gives me cause to introduce certain modi- 
fications or to suggest some alternatives. There- 
fore Table 2 is offered as a new key to the system, 
and Fig. 1 shows how the symbols (now simpli- 
fied) can be plotted on squared paper (8 squares 
high, 25 squares long). 

These modifications improve the system in 
a number of ways. 

Life-form. Inasmuch as the former "trees" 
and "shrubs" did not necessarily refer to plants 
at the adult stage of their development but only 
to woody individuals of a certain height, at a 
certain time, it is best to designate by a single 
symbol all such individuals. This leaves category 1 
with only 5 instead of six alternatives. It will 
also be noted that the symbols now proposed 
for lianas (L) and epiphytes (E) are much easier 
to draw. 

Stratification. In substituting this heading for 
"size," a more objective picture is conveyed, 
and the sliding scale principle is better applied. 
Thus, all symbols followed by the same figure 
(1 to 7) will situate that life-form at the same 
level (e.g. in the same layer) as all other life-forms 
coupled to the same figure. This was not so 
previously: e.g., Tt and Ht were 23 metres apart. 
Now, the synusiae are well separated, and the 
layers are united! 

Figure 2 shows some types of vegetation that 
had been previously plotted with the 1951 key. 
Although this may be an improvement in practical 
procedure, it still does not allow to show a certain 
number of features (see 7, Table 8), which, for 
certain purposes, it may be desired to illustrated. 

Coverage. Therefore, some of these features 



are added here and shown in Figs. 3, 4, and 5. 
The two diagrams previously published (7, Figure 
1 3) and now re-plotted show in their upper layer 
a certain total crown coverage that could very 
well be the result of different stem growths. 
Inasmuch as the system so far has made little 
provision for emphasizing the closeness of stems, 
a new means is now offered to show this. Figs. 
3, 4, and 5 outline two different approaches: one 
by direct drawing of independent stems and 
crowns (and modification of the crown symbols 
from circles to ellipses) and the other by drawing 
lines that show how many times closer the stems 
are than the normal coverage symbol allows for. 

A further refinement can be introduced to 
represent the principal outlines of crowns as 
indicated in Fig. 6. There would be no particular 
merit in extending such categories to herbs, lianas, 
epiphytes or bryoids, inasmuch as leaf type 
pretty well determines outline anyway. 

Of course there always remains another alter- 
native, which would consist in prolonging the 
strip many times: 50, 75, 100 or more squares, 
instead of 25, would allow an almost literal repre- 
sentation of the principal life-form types along this 
linear transect. 

Function. Relative deciduousness (or leafing 
periodicity) is of course quite important for it 
affects volume at different seasons. But there is 
another item of great importance to vegetation 
analysis, the dispersal function. This has been 
discussed at some length by Dansereau and Lems 
(6) . It is very remarkable that the different layers 
show such marked contrasts in the distribution 
of, for instance, wind- and animal-dispersed 

In a recent paper, Keay (8) analyzes a second- 
growth forest in Nigeria where he finds that wind- 
disseminated tree species are dominant. On the 
contrary, in the Canary Island laurel forest, 
bird-dissemination is the rule at the tree level. 
In the temperate forests of the Pacific, a very large 
number of trees are dispersed by wind: from 
Alaska to northern California, the tallest members 
of the needle-leaved forest (Picea, Tsuga, Sequoia, 
Pinus) have very light diaspores, whereas a greater 
number of the shrubs and herbs have fleshy 
fruits. The Mediterranean area of California, 
on the other hand, has a vast array of oaks and 
some fleshy-fruited trees like Arbutus. A similar 
situation obtains in New Zealand, with the 
northern forest rich in fleshy fruits, the southern 
rich in nuts and small winged diaspores. 


Although this system was devised essentially, 
in fact exclusively, in order to show the features 
of vegetation and later and otherwise relate them 
to environmental elements (causal or not), in 
actual recording it is found useful to plot a certain 
number of the latter, for instance site conditions. 
Table 3 offers a small repertory of the usual 
alternatives: climate, relief, qualities of soil, 
land use. 


The problems of mapping are very numerous 
and each scale presents its own difficulties. No 
easy rules can be established (and surely none are 
explicitly recognized) when it comes to stripping 
a large-scale map (say 1 inch = 1 mile) of its 
detail if the information is to be relayed to a 
smaller scale (for instance, 1 inch = 8 miles). As 
a matter of fact, it is rarely a matter of omitting 
detail; it is more likely a matter of interpreting 
the units of the large-scale map so as to lump 
them into units of a greater order of magnitude. 
This involves a judgment, something which is not 
strictly a matter of record or observation, of 
compilation of features that already lie on the 
map itself, but something that is essentially classi- 
ficatory. An example of this is provided by the 
excellent maps of the Toulouse group, for instance 
Gaussen's (J) Perpignan sheet. The colours 
represent each a "series" (we would say a sere): 
a number of plant communities, usually of in- 
creasingly complex structure, which lead to a 
terminal stage. 1 The intensity of the colour, on 
the other hand, does indicate changes of structure. 
This procedure appears to me worthy of standard- 
ization and lends itself to a very useful procedure. 
Thus, for instance, the kauri forest area of New 
Zealand or the highland pine region of the Island 
of Luzon (Philippines) in their developmental 
stages from grassland or scrub to mature forest 
could be represented by a series of structure 
diagrams, and on a map by the corresponding 

Schmid (9) had already recognized the desir- 
ability of producing structure diagrams in the 
margin of vegetation maps in order to show the 
range of variation which must of all necessity be 
involved in any one category of vegetation type 
[except on a very large-scale map which really 
attempts to show the exact status of the plant- 

l This is generally the climatic climax although occasionally an edaphic or topographic climax. See ( 3) for a fuller discussion 
of this topic. 



cover (Pflanzendecke) at one time!]. 

It is hoped that the present schemes lend them- 
selves to fairly wide application and can be used 
to reveal at a glance the many-sided aspects of 
vegetation types that play an important role in 
the landscape. It may well remain to demonstrate 
to the biologist and, to the paleontologist what 
the historical and physiological conditioning of 

these types really is. It is granted, of course, that 
such considerations are fundamental, but the fact 
that they are virtually disregarded in the present 
instance, should not disguise the urgency of a 
better comparative understanding of the 
functional aspect and distribution of vegetation 
types as they are to be observed in the living 
landscape at present. 


(1) Dansereau, Pierre, 1951, Description and 

recording of vegetation upon a structural 
basis, Ecology, 32 (2): 172-229. 

(2) Dansereau, Pierre, 1953, Structural units 

of vegetation in tropical and temperate 
climates with special reference to Pacific 
areas, Proc. Seventh Pac. Sci. Congr., 

(3) Dansereau, Pierre, 1956, Le regime clima- 

tique regional de la vegetation et les 
controlcs edaphiques, Rev. Canad. Bio/., 

(4) Dansereau, Pierre, 1957a, Biogcography : An 

ecological perspective. The Ronald 
Press Co., New York, xii + 394 pp. 

(5) Dansereau, Pierre, 1957b, A preliminary 

note on the structure variations of 

temperate rain forest, Proc. Eighth Pac. 
Sci. Congr., 4 (Botany): 407-436. 

(6) Dansereau, Pierre and Lems, Kornelius, 

1957, The grading of dispersal types in 
plant communities and their ecological 
significance, Contrib. Jnst. Rot. Univ. 
Montreal, 71 (in press). 

(7) Gaussen, H., 1946, Feuille No. 78: Perpig- 

nan. In: Carte de la Vegetation de la 
France (1 :250,000). Centre National de 
la Recherche Scientifique, Paris. 

(8) Keay, R.W.J., 1957, Wind dispersed species 

in a Nigerian forest, Journ. EcoL, 45(2): 

(9) Schmid, Emil, 1954, Anleitung zu Veget- 

ationsaufnahmen. Vierteljahrsschrift 
der Naturforschenden Gesellschaft in 
Zurich, 1C (1954), 1,37pp. 


Six categories of criteria to be applied 




















Table 1. 


2. SIZE 

t tall (T: minimum 25 m) 

(F: 2-8 m) 
(H: minimum 2 m) 

m medium (T: 10-25 m) 
(F,H: 0.5-2 m) 
(M: minimum 10 cm) 

1 low (T: 8-10 m) 

(F,H: maximum 50 cm) 
(M: maximum 10 cm) 


b barren or very sparse 

i discontinuous 

p in tufts or groups 

c continuous 


I I deciduous 
1 1 1 1 semideciduous 
"|| 1 1 evergreen 

evergreen-succulent ; 
or evergreen-leafless 

n <^> needle or spine 
g A graminoid 

medium or smal 




P O thalloid 


succulent; or fungoid 



Table 2. 

A revised scheme of the six categories of criteria to be applied to a structural description ol 
vegetation types. 










erect woody plants 

climbing or decumbent woody s 1 



e : 



j j 




1 more than 25 metres 

2 10-25 metres 

3 8- 10 metres 

4 2-8 metres 

5 0.50-2 metres 

6 0.10-0.50 metres 

7 0.0-0. 10 metres 


b barren or very sparse 

i interrupted, discontinuous 

p in patches, tufts, clumps 

c continuous 





evergreen- succulent ; 
or evergreen-leafless 



v V 

P o 

needle or spine 


medium or small 








| | membranous 
succulent; or fungoid 









Table 3. 
Categories and symbols for recording site conditions. 









(A, B, C horizons) 

bedrock | V V y" 















use symbols of compass, 
N, S, E, W 

e.g.: NW 


use Koppen symbols 
e.g.: Dfc 




r' C 

f C 




? ? c 



Wl W2 W3 


W5 W6 W7 







A E3 



L5 CD 


A E6 





H4 H5 H6 H7 M6 M7 

Fig. 1. A graphic representation of all the symbols combining criteria 1 and 2 of Table 2. This is a new scheme, super- 
seding the one proposed in 1951. 




Aceretum rubri Wldhze(ozb) WZdhzi W3dhzp W4dhzi 

WGdhzi(azb) HGdhzb W7dazb H7dazb 

Betuletum populifoliae W2dazi W3do(h)zb W4dhzb 

W6dazb H6dvzb H7dg(o,v)zp 


Fig. 2. A red maple stand and a wire-birch stand plotted according to the new scheme (Table 2 and Fig. I). (Compare 
with Figs. 13 and 14 of the 1951 version.) 




Fig 3. The upper la>cr of the red maple stand (see Fig. 2), showing different distribution of stem and crown spaces cor- 
responding to identical coverage and illustrating two ways in which the stand can be plotted. 





000000 000000 00 00 

Fig. 4. The upper layer of the wire-birch stand of Fig. 2, illustrating variations in stem within an identical total crown 
coverage and two ways in which these differences can be plotted. 





Fig. 5. Another variant of crown-stem distribution in a wire-birch stand (see Figs. 2 and 4). 




Fig. 6. A series of crown outlines for tall woody types (W 1, 2, 3), which can fit the perimeter of the symbols in Fig. 1. 





Department of Biology, Kyushu University ', Fukuoka> Japan. 


It is possible to say that there may be consider- 
able subjectivity in classifying vegetation, even if 
we adopt the abstract method of Ziirich-Montpel- 
lier's school, attaching great importance to 
characteristic species, which is commonly em- 
ployed in continental Europe. We have a strong 
urge to find some more objective method of 
classifying vegetation. Fortunately, our attention 
was called to Gpodall's objective method (I) 
for the classification of vegetation. This method 
is based on positive interspecific correlation of 
major component species by making use of the 
values of their frequency. We modified his 
method to some degree (I), and used it and 
Sorensen's method (5) for classifying epiphyte 
vegetation growing on beech trees in Mt. Hiko, 
Southwest Japan. 


The concept on which the modified method of 
classifying vegetation is founded is that if there is 
neither species of positive nor of negative inter- 
Specific correlation of significance level in any 
given plant group, such a group is conceived of 
as being of homogeneous construction. Then, 
the epiphyte vegetation is classified objectively 
by four procedures according to this method of 
consideration, practically almost the same as 
Goodall's methods (I) of separating groups. 

(a) Procedure I. The species of higher fre- 
quency, as well as of positive interspecific correla- 
tion of P< 0.001 level of significance were used 
for separating groups by excluding all quadrats 
in which the single correlated species did not 
occur, until this particular species was no longer 
present in a given group. One of the final groups 
of residue was considered in its possible combina- 
tion with another group, if there was not recog- 
nized any species of positive interspecific correla- 
tion which reached a high significance level in a 
newly combined group. We expect, however, 
that there may exist some species of negative 
interspecific correlation of P< 0.001 or 0.0 1< 
P< 0.001 level of significance even in such a final 

group conceived to be certainly homogeneous as 
judged by positive interspecific correlations of 
significance level. 

(b) Procedure II. The species of lower fre- 
quency and simultaneously of positive inter- 
specific correlation of a high significance level 
which is quite the same as Procedure I, were used 
for separating groups by excluding all quadrats in 
which the single correlated species occurred, as 
well as by excluding those in which it did not 
occur (Procedure I). Recombination of the final 
groups of residue was made by the same method 
as Procedure I. 

(c) Procedure III. As a substitute for positive 
interspecific correlation, the species of higher 
frequency and of negative interspecific correla- 
tion of high significance level were used for sepa- 
rating groups like Procedure I. In this case too, 
we may expect some species of positive inter- 
specific correlation of the same level of signific- 
ance remaining in the final groups. 

(d) Procedure IV. As a substitute for the 
species of higher frequency, we used those of 
lower ones for separating groups. The treatment 
is quite the same as Procedure II. 

Procedure I is clearly the best one for classifying 
the vegetation of epiphytes, as shown in Table 1 , 
notwithstanding the difference that exists among 
the three series of methods different in quadrat 
size and number found in random sampling on 
five beech trees: one standing at the peak, Kita- 
dake, of 1,150 m alt.; two at the ridges of 925 m 
and 930 m alt. ; and the other two at the moun- 
tain-slopes of 910 m and 960 m alt. 

The smaller the quadrat size, the more numer- 
ous the groups become. Fig. 1 shows the simi- 
larity of correlation among all the groups recog- 
nized in the three series of groupings by means of 
the sampling data of the different kinds of quadrat 
size. Each group is designated by the names of 
two important species which bear the highest and 
the next best figures of positive indicator value. 
However, if there are only a very few or not any 
species having positive indicator values of signi- 
ficance level in a group, we indicate it by enclosing 
the name of the species not reaching significance 

t Presented by F.E. Egler. 

Table 1. 

Success of different procedures in dividing data, 
which were obtained from different sampling 
sizes of quadrats, into homogeneous groups. 






No. of interspecific 






with P* 







< 0.001 















































1 f 
















* The table shows the number of Positive (in Procedures III & IV) 
and negative (in Procedures I & 1J) interspecific correlation of sig- 
nificance level in the final groups. 

level in parenthesis within a rectangular frame of 
broken line. The degree of independence of every 
epiphyte-group classified in these ways is ex- 
pressed by the species number of positive (in 
Roman type) and negative (in italics) indicator 
values (1, 2) of significance level (P < 0.01), 
which are shown beneath the rectangular frame 
(Fig. 1) in which is written the group name. 
The similarity of correlation among the groups, 
depending on the three simultaneous series, is 


expressed by using the "Quotient of Similarity" 
(=QS, 5). In Fig. 1, the highest value of this 
Quotient between one group of one series and a 
group of the other series is shown near the con- 
nected line between groups, and the correlations 
between them are shown by various forms of 
lines. Only the major connection-lines of simi- 
larity, excluding minor ones, are shown. In 
comparing groups between two series with each 
other, to which both groups belong, the thick line 
indicates the closest correlation where the highest 
similarity is recognized between the two groups. 
The thick broken~lines having a triangular arrow- 
head indicate that a given group from which the 
arrow-headed broken line started has the highest 
value of QS correlating with the group indicated 
within those of the other series. According to QS, 
especially the groups centering around those 
having the highest values of QS, as shown in 
Fig. 1, we may regroup the epiphyte groups of 
each series into four group-types. 

Of every group in the series (Series 1) which 
resulted from the Procedure I of arranging the 
data obtained from the smallest size of quadrat 
(500 sq cm each), Table 2 shows the ecological 
distribution concerning vertical position and expo- 
sure on trees and topographical positions of host 
trees. The state of ecological distribution of 
every group in the other two series, both Series 2 
and 3 obtained from Procedure I, which resulted 
from the data of larger quadrat (1,000 sq cm 
Series 2, and 1,500 sq cm Series 3) is not so 
different from that of the smallest sampling series 
(Series 1) shown in Table 2. 

As shown in Fig. 1, in each series, several 
somewhat homogeneous groups were recognized 
as the result of objective classification of epiphyte 
vegetation in terms of interspecific correlation, 
while there were some groups which had few 
species of positive indicator value of significance 
level. In order to make clear the correlational 
similarity of floristic composition among the 
independent groups of each separate series, 
Sorensen's grouping method (5) based on the 
value of QS was performed. An outline of this 
method is shown in Fig. 2 and Table 3. For 
example, we will explain the method of integra- 
tion of those epiphyte groups in the Series 1 de- 
rived from sampling data of the smallest quadrat 
(See Table 3 and Fig. 2). The order of arrange- 
ment of the epiphyte groups was the same as that 
(Series 1 ) of Fig. 1 . At the level of QS = 40 (Table 
3 and Fig. 2), the Homalia japonica Homalio- 
dendron scalpellifolium group (Group 1) and the 
Thuidium cymbifolium - Homaliodendron scalpel- 



Series 5 

Series 2 

Scries I 

Series 3 

~ ~~ ^ * - "~ "* 

_ 3 3 -^-\ 

Fig. 1. Classificatory schema of epiphyte communities, depending on Procedure I on the basis of the data obtained from 
the three kinds of sampling sizes of quadrats (Series 1 means the grouping resulted from 423 quadrats of 500 sq cm; Series 2, 
that of 282 quadrats of 1,000 sq cm; and Series 3, that of 150 quadrats of 1,500 sq cm). The numerical figure written in 
Roman type beneath every rectangular frame in which is shown the group name, indicates the number of species which 
has the positive indicator value of significance level in that corresponding group, and that of italic type is of negative indicator 
value. The numerical figures situated near a connected line indicate the value of Quotient of Similarity between the groups 
connected with the line. 

lifolium group (Group 5) were unified (QS = 42) 
into one group (Group 2), and at the same time 
Groups 3 and 4 (QS = 41) were combined into one 
group (Group c) ; at the QS = 30 level, the Group b 
and e (QS = 32) into the Group B, and the Group c 
and d (QS = 33) into the Group C; and so on. 
The number of groups differs to some degree at 


different levels. Noting Fig. 1 and Table 3, we may 
say that the group which has only a few species 
bearing indicator value of significance level among 
the component species, correlates with the other 
groups by the lower value of QS. Though the 
number of groups differ much in the different 
series which differ from each other by the size of 


Table 2. 


Showing the state of the ecological distribution of epiphyte groups by the numbers of quadrats. 
Vertical distribution on trees: trunk-bases (Tb), trunks (T), lower parts of crowns (Cb), interior parts 
of crowns (C), and top-most parts of crowns (Ct). The distributed side on trees: open side (O), 
intermediate side (M), and sheltered side (S). Distribution related to topography, viz., topographical 
distribution of host trees on which the epiphytes grow: a beech tree standing at the peak, Kitadake, 
of 1,150 m (Fj), at the ridge of 930 m (F 2 ) and 920 m (F 3 ), and at the slope of 910 m (F 4 ) and 960 m (F 5 ) 
alt. The figures having a thick underline indicate the predominant position of a given group. 






Group ^x. 

Vertical distribution 
on trees 

on trers 

related to 









F 2 





Parmelia loeirior 
Uhtd uispula 












Diiranolpmd tlagiliformc 
MetzQerid conjugate 








1 1 






Graphic sp. 
Cetrdsia colldta 














Bouldyd mitteii 
An z id. japonic a 












1 1 

Ortiwdicranum kaktodeose 
Bouldyd mite nii 












Ptirmelid homogcries 
Frulldiud, monilidta 














AnomotLon girdkUi 
DolifomHm cymbffolid 












1 1 

forsst/vcmid ( ryptuxvides 
Porclld irentisia 

















fhuidium. {y/nbifol/w* 
HomdiodtAomn siidpeJIijolium 







homalid idpo/uca 
Hoindliodendnon <4djpcV(folium 













Total no. of quadrat 












quadrat for drawing data from the epiphyte 
vegetation, the larger the size of the quadrat, the 
fewer the number of groups becomes. Notwith- 
standing the difference of series, however, in any 
case the epiphyte vegetation is classified roughly 
into two groups (e.g., Fig. 2): one is of epiphytes 
growing on trunks and trunk-bases, and the other 
those of the interior parts of crowns and the top- 
most parts of crowns. 

At a certain level where we can determine the 
value of QS, as we hoped, as a given criterion of 

integration of epiphyte groups, we can unify 
those groups into the higher rank of group at a 
given level, as well as we can understand the 
degree of similarity of floristic composition among 
those groups. Moreover, it is certainly note- 
worthy that the order of arrangement of those 
groups (Figs. 1 and 2) which are naturally 
arranged in the case of classifying vegetation, 
such as their ordering positions from the top- 
most parts of crowns down to the trunk-bases, 
is quite in accord with the environmental gradient. 



Group QQ 

U/ot<* crisp. 

Dicntnolom* f. - 
Metzgeri* nnj. 

Grttphif //>. - 
Cetrttrk* coi. 



Boulaya. mitt - 

Btuya mitt. 

k - 

c JhuMium cymk - 
i Horn* lie* Jc 





Fig. 2. Series 1 of Sorensen's grouping method, in which the integration was performed at the foundation of final groups 
which were recognized in terms of classifying the epiphyte vegetation through Procedure 1 on the basis of the data obtained 
from 423 quadrats of 500 sq cm. 




In order to compare those groups resulting from 
such methods of treatment with the five epilias 
(3) of epiphyte communities which we have 
studied and designated to the same epiphyte 
vegetation in the beech forests of Mt. Hiko (4), 
and moreover to make clear the correlation be- 
tween those five epilias and the groups based on 
the present treatment, we selected six correspond- 
ing epiphyte-groups (the Group A, B, C, D, E, 


and F in Figs. 2 and 3) at the level of QS = 32 
(Fig. 2) in the series of sampling with the smallest 
size of quadrat (Series 1). Comparison was made 
between them by using the values of QS of each 
one (Fig. 3). Here the correlational similarities 
among those groups are clearly recognized. 
Indication of the similarity between an epilia and 
a group, which is shown by the lines and the 
numerical figures of QS, was done in the same 
way as in Fig. 1. What is written beneath each 
epilia-name surrounded with a rectangular frame 
is the number of the characteristic species, and 
similarly that of the epiphyte-group name means 


Table 3~~~~ 
Showing the values of QS between the epiphyte groups at four different levels in Series 1, which 

is sh 





in Fig. 2. 

d. b. e. d. c. g. f. h. 
























































































10. 8. 8. 4. 3. 6. V. 2. 6. 1. 


F. D. E C. B. A. 

the species number of the positive (in roman type) 
and negative (italic) indicator value of significance 
level. From these figures and the connected line, 
the correlational correspondence between each 
epilia and group is comprehensible to us. It is 
possible that the epilia or group which has corre- 
spondingly fewer characteristic species or fewer 
species bearing significant indicator value among 
the component species, in either case, does not 
necessarily show any correlational correspondence 
between such epilia and group. 

Accordingly, because of the success of our 
several years of varied studies of corticolous 
vegetation in the beech forests of Mt. Hiko, South- 
west Japan, we would draw the conclusion that the 
following four distinct communities of epiphytes, 
which are considerably developed there can be 
recognized: the Thuidium cymbifolium-Homalio- 
dendron scalpellifolium epilia (3) or Thuidium 
cymbifolium-Homaliodendron scalpellifolium group 
(at the lower part of tree trunks), the Pterobryum 
arbuscula - Anomodon giraldii epilia or Forsstroe- 

mia cryphaeoides - Anomodon giraldii group (on 
trunks), the Cetraria collata f. nuda - Boulaya 
mittenii epilia or Boulaya mittenii - Frullania 
moniliata group (at the upper part of trunks and 
on boughs in crowns), and the Ulota crispula- 
Pertusaria sp 2 epilia or Parmclia laevior - Ulota 
crispula group (at the top-most part of the crowns). 
It will be probable that the other three remains 
shown in Fig. 3, viz., the Graphis sp t - Pertusaria 
spj epilia, Graphis sp. -Cetraria collata group, and 
Dicranoloma fragiliforme - Metzgeria conjugata 
group, are all unlikely to be developed into 
independent social units or groups. 


In cases where it is difficult to distinguish plant 
community individuality, as in corticolous com- 
munities consisting chiefly of bryophytes and 
lichens, it seems to us to be possible to make a 
classification of such vegetation and to systema- 
tize the communities objectively. This would be 



e U/ota crispu/a - 
* Pcrtusctria sp.2 



Epiphyte group 

* Bou/dydL mittenii 



Perl us Aria sp. i 

Ptervbryum drbuscula - 
Anomoaon gimldii 


r\ Pcirmelici laevior - 
U/ota crisputa 

** fragl/iforme - 


sp. - 

.'y p Bou/ayd jnittenii - 
' Frvllajrii* noni/idtd 


D Forsstroenu'ci cryphdeoides - 
u> Anomodon girctldii 

L Uuidium cymtifolium - Y ^\\ ~n' !f * l " m rvmtifnlium - 
' homaliodendmn scalptHifolium \ 54 541 ' flomaliot/ein/ron stxtlpellifolium 

5 (9 

Fig. 3 Classificatory schema of epiphyte communities, showing so as to make comparison between each epilia and group 
by means of the value of QS. Indication of correlational similarity between them is the same to Fig. 1. 

done on the basis of floristic composition, by 
classifying the communities into some homoge- 
neous groups by making use of the value of inter- 
specific correlation based on the frequency of 
major component species and in the light of the 
indicator value of significance level (I, 2). These 
groups would then be integrated into an appro- 
priate number of groups of higher rank at a 
wanted level in accordance with the value of 
"Quotient of Similarity" (-QS, 5) which is based 
on the frequency of component species. 


(1) Goodall, D.W., 1953, Objective methods for 

the classification of vegetation, Austral. 
Jour. Bot., 1: 39-63, and 434-456. 

(2) Hosokawa, T., 1955-56, An introduction of 

2x2 table methods into the studies of the 
structure of plant communities, Jap. 
J. Ecol., 5: 58-62 (1955), 93-100 (1956), 
and 150- 153 (1956). 

(3) Hosokawa, T., Omura, M. and Nishihara, 

Y., 1954, Social units of epiphyte com- 
munities in forests, VIHe Congr. Intern. 
Bot. Paris, 1954. Rapports comm. Sect. 
7: 11-16. 

(4) Omura, M., Nishihara, Y. and Hosokawa, 

T., 1955, On the epiphyte communities 
in beech forests of Mt. Hiko in Japan. 
Rev. Bryol. Lichen. 24: 59-68. 

(5) Sorensen, Th., 1948, A method of establish- 

ing groups of equal amplitude in plant 
sociology based on similarity of species 
content. Dei Kong. Dansk. Vid. Sel. 
Biol.Skr.5(4): 1-34. 




University of Kansas, Lawrence \ Kansas, U.S.A. 

Let me begin with a brief report on what has 
been accomplished so far in mapping the vege- 
tation of the Pacific Region. As you can imagine, 
the accomplishments are not at all uniform, but 
they are nevertheless very promising. There are, 
of course, numerous vegetation maps of very 
small areas. I shall ignore these here and report 
to you only on maps of countries. 

In North America, a vegetation map of British 
Columbia was published recently, and maps of 
Mexico and Guatemala have been available for 
some time. A map of Alaska is now in press, and 
there has been a small map of the United States 
for a long time; a larger one is now in preparation. 
Most of the Central American states remain to 
be done. This is a relatively favourable report, 
especially if we include the work now in progress. 

In South America, conditions are much less 
satisfactory, and Peru is the only country of which 
a reasonably detailed vegetation map exists. 

The most favourable report comes from Austra- 
lia. That continent has now been mapped, and 
the new vegetation map by Williams has been 
published recently. New Zealand has also been 
mapped, as well as New Guinea. On the other 
hand, the various Pacific islands have been 
mapped very unevenly, if at all. 

In Asia, progress varies from one country to 
the next. The new Soviet map covers a large part 
of the Asian continent. Indonesia, the Philip- 
pines, and Taiwan have vegetation maps that need 
some further work. There are good but small 
maps of Korea, Manchuria, Thailand, and Burma. 
The greatest gaps are China and Japan, Malaya, 
and North and South Vietnam. 

Actually, vegetation maps are now available 
for the entire Pacific Region, and the International 
Bibliography of Vegetation Maps which I hope 
to publish some time soon contains over 200 
references to countries bordering on the Pacific 
Ocean. The scale and quality of these maps vary 
within wide limits, and many of the vegetation 
maps are too small or too generalized to be of 
much use. But modern vegetation maps can be 
useful instruments in a variety of scientific investi- 
gations, as well as in planning and managing the 
use of our lands. This has been demonstrated 
many times. 

If, therefore, it is evident that vegetation maps 
are useful, I should like you to consider the possi- 
bility of preparing a vegetation map of your 
home land if a good one at a large scale does not 
already exist. If you are not in the position to 
prepare such a vegetation map, you can perhaps 
encourage one of your colleagues to do so. 

The maps should be useful to as many people 
as possible and not just to those of one country. 
For this reason, a certain degree of coordination 
is valuable. In addition to suggesting that new 
and large scale vegetation maps be made of every 
country of the Pacific Region, I should therefore 
like to propose that the vegetation be mapped in 
its major physiognomic and structural features, 
and that these broader types be refined according 
to their florist ic character by listing the dominant 
genera or species wherever that is feasible. 
Structure and floristic composition are the two 
basic features which all vegetation types on earth 
have in common. They should therefore be given 
priority on vegetation maps. If other features 
are to be added, such as ecological aspects of the 
habitat, they might well be in the nature of a 
supplement rather than primary information. 
Such supplementary information may be given 
wherever this is feasible and useful, but I think 
it should be only in addition to the two basic 
features of physiognomy and floristic com- 
positon, and not instead of them. 

Of course, there are many ways to make a 
vegetation map, and it is quite possible that 
authors may wish to adapt their maps to the 
circumstances of their respective countries. The 
possibilities of variation are greater than many 
authors realize; and while a vegetation map 
should not be cluttered, there is, nevertheless, no 
need to ignore any chances to make the map 
content more meaningful. 

Those who wish to make large scale maps will 
find interesting and stimulating models in Carl 
Troll's vegetation map of the Nanga Parbat area 
in the western Himalaya, or in Heinz Ellenberg's 
vegetation-site map of Leonberg, Germany; both 
maps are at the scale of i : 50,000. Larger scale 
phytosociological maps, as published under the 
direction of Louis Emberger or Reinhpld Tiixen, 
are perhaps not yet feasible for the Pacific Region 



as they require more detailed information than 
is generally available. As far as I know, only 
some Japanese scientists have successfully at- 
tempted to prepare such maps, and only in very 
limited numbers. 

Inspiration for small scale maps may be gained 
by studying Kurt Hueck's map of Germany 
or Henri Gaussen's map of France, both at 1 : 
1,000,000. More recently, our Soviet colleagues 
have published vegetation maps of their country 
at 1 :4,000,000, and Williams published his new 
map of Australia at 1 :6,000,000. My own map of 
the United States is at 1:14,000,000, and this 
small scale is justified only because the map was 
included in an atlas. 

The preparation of a new vegetation map of 
the United States at a larger scale is now in pro- 
gress, but I do not expect to complete the manu- 
script for another three or four years. For the 
large countries of Australia, Canada, China, the 
Soviet Union, and the United States, scales of 
1 :4,000,000 or 1 :6,000,000 can be justified, but 
for smaller nations a scale of 1 : 1,000,000 should 
be the minimum. 

1 mentioned that many vegetation maps of 
the Pacific Region are of too small a scale or too 
generalized to be very useful. This need not be 
discouraging because the field of vegetation map- 
ping is developing only now. The scale can always 
be enlarged and more details can be added. Far 
more significant are the map content, its organi- 
zation, and representation. A consideration of 
these features is fundamental and, indeed, im- 
perative before preparing a vegetation map. 

Today I should like to present to you two 
problems for discussion. The first one is that of 
the map content. Some authors argue that a 
vegetation map should contain nothing but 
information on vegetation; others want to show 
more, especially ecological features and also land- 
use data. A strong case can be made for both 
sides. I do not propose here that a decision should 
be taken in support of one approach to the ex- 
clusion of the other. Both approaches are valu- 
able and justifiable. But I do feel that the matter 
should be discussed now so that those who 
return home to prepare or promote a vegetation 
map of their area will see more clearly what 
problems they will have to face. 

The argument for showing only vegetation on 
a vegetation map is strong and logical. Vegetation 
is so complex that the map can always be filled 
with as much detail as the scale permits. If the 
over-all character of the vegetation is simple, 
then further detail can be shown by introducing 


various structural aspects of the vegetation, 
differences in the lowest synusia, floristic varia- 
tions, transitions associations, sub-associations, 
variants, sub-variants, and other features. The 
map is then filled to capacity, and the entire 
information refers only to vegetation. By exclud- 
ing all other kinds of information, the vegetation 
can be described in the greatest possible detail. 

But there are others who feel that some detail 
of the vegetation can or should be sacrificed in 
order to introduce ecological information. There 
is, of course, no doubt that such information is 
revealing and valuable, and therefore should be 

However, the introduction of non-vegetational 
data creates a conflict: what part of the vegeta- 
tional information is to be sacrificed for the sake 
of ecological data, and also, how much ecological 
information may be introduced without under- 
mining the character of the map as a vegetation 
map? A further problem is the kind of ecological 
information to be shown in a vegetation map. 
Should it be any kind, whatever the author 
happens to favor at a particular time and place, 
or should it be so organized that a logical system 
can be applied? If so, what specific ecological 
data are to be selected as it is impossible to in- 
dicate the environment comprehensively? 

Usually, vegetation maps with ecological 
information lack unity, but they are nevertheless 
very numerous. This shows that many authors 
place much value on this approach. On the other 
hand, there are only very few vegetation maps 
that are based on a strictly organized system 
which includes both vegetational and environ- 
mental features. I am thinking of the works by 
Heinz Ellenberg and Henri Gaussen. There are 
also vegetation maps which are strictly vegeta- 
tional but which have strong and definite ecolog- 
ical implications. The maps by Reinhold Tiixen 
and his collaborators belong in this class. On the 
maps by Hueck, or by Lavrenko and Sochava, 
ecological information is sometimes given, some- 
times implied, and sometimes omitted. Carl 
Troll fits the vegetation into the landscape, so 
that his vegetation map, too, is ecological in 
character; his method is most enlightening but 
is most readily applied to mountainous terrain 
on large scale maps. 

The fact that the most interesting or the most 
valuable vegetation maps were not made in the 
Pacific Region is not at all a cause of concern. 
What really counts is the basic ideas on which a 
vegetation map rests. Once grasped, these ideas 
can be applied in the Pacific Region as well as 



anywhere else. The problem of the map content 
is well worth discussing because the character 
and the usefulness of the maps depend on it. 

The other feature I wish to present to you for 
discussion is the use of color. Several authors 
have shown that the judicious application of 
color and color patterns can greatly enrich the 
map content. Many years ago, Riibel in Swit- 
zerland proposed a color scale for Swiss vegeta- 
tion maps. This scheme turned out to be too 
limited and was not often applied. Much later, 
Emil Schmid, also of Switzerland, was careful to 
select harmonizing colors to facilitate map reading 
but refrained from proposing definite colors for 
given vegetation types as Riibel had done. 
Schmid has made more of a contribution than it 
seems because many vegetation maps have dis- 
turbing color schemes. Clashing colors make 
map reading difficult and easily suppress im- 
portant features on the map. 

Sochava of the Soviet Union briefly discusses 
the use of color on vegetation maps and proposes 
that the color scale used on the Soviet map be 
employed elsewhere, too. But he failed to indicate 
just how he arrived at his color selections, and 
therefore it is not possible to use his ideas sys- 
tematically in other countries. 

The only person who so far has presented a 
complete and logical color system for use on 
vegetation maps is Henri Gaussen of France. 
He systematically uses the sequence of colors in 
the spectrum and different patterns within each 
color to express vegctational and ecological 
information. The system has been criticized by 
Sochava, and indeed, some modifications may be 
desirable. But the fact remains that so far, and 

as a method of using colors on vegetation maps, 
Gaussen's work is unique. 

If you agree with me that vegetation maps 
should be made of the entire Pacific Region, then 
we should ask ourselves whether or not colors 
should be used uniformly throughout the region 
in order to tie the area together and integrate the 
individual maps into a larger whole. Question 1 
is therefore: should a color scheme be prepared 
for the vegetation of the Pacific Region, or should 
the use of color be left to the discretion of each 
author? If a more or less uniform system seems 
desirable, then there follows question 2: should 
the use of colors be restricted to vegctational 
information, or may colors refer to ecological 
data as well? The more systematic the use of 
color, the more readily can the reader distinguish 
the major groups of features at a glance. 

Finally, there is the use of symbols which may 
be employed to enrich the vegetational and/or 
ecological information. And again, as in the case 
of colors, should symbols be used indiscrimi- 
nately or should they be restricted to any one 
type of features? Uniformity of approach cer- 
tainly facilitates the appreciation of vegetation 
maps, but whether such a form of coordination 
can or should be achieved is a matter for dis- 
cussion here and now. 

Certainly the present state of vegetation map- 
ping in the Pacific Region leaves much to be 
desired. Certainly, enough vegetation maps 
have been published all over the world that the 
basic ideas can be grasped and developed. 
I want to urge you to clarify the thinking of all 
of us who arc interested in preparing vegetation 
maps of the Pacific Region. 





Museum National d'Histoire Naturelle, Paris, France. 

People who deal with the subject of the vege- 
tation of the islands of the Pacific always em- 
phasize the youthful nature and the ubiquitous 
and more or less ruderal flora of the savannahs 
of these islands. 

The "savannahs" are qualified as "secondary" 
and as being the outcome, in fairly recent times, 
of the clearing of forests for cultivation and the 
subsequent abandonment of the cultivated areas. 

The floristic lists given are convincing in this 
respect, apart from some reservations which 
should be made. 

Virot (1957) considers that the "savannahs" of 
New Caledonia are of secondary origin and the 
direct outcome of the deforestation of the island. 
The first explorers already pointed out the pre- 
sence of vast "savannahs." However, they 
represented only "a degraded and secondary 
aspect of the vegetation and . . . they are made up 
almost entirely of the species introduced, parti- 
cularly the graminaceous species." (Virot, p. 
62). "The secondary "savannahs," existing at the 
time when France took possession of New Cale- 
donia, were, from the beginning, used in part as 
natural pasture land. Generally speaking, they 
consisted of stretches of dry grasses, among 
which the ubiquitous pan-tropical plant pre- 
dominated, the graminaceous species for the 
most part, of which the principal ones are Hetero- 
pogon contortus Roem., Aristida pilosa Labill., 
Eleusine indica Gaertn., Dactyloctenium aegyp- 
tiacum Willd., etc..." (Virot, pp. 65 and 66). 
"First of all, it is all but certain that almost all 
the grassy stretches of New Caledonia have no 
other origin than that of the destruction of virgin 
forest by fire." (Virot, p. 68). "Among the 
graminaceous species may be mentioned, among 

others: Heteropogon contortus Roem. and Schult; 
pan-tropical pyrophytic plant, typical of "se- 
condary savannahs", and Themeda triandra 
Forsk; which are, further, widely prevalent in 
the interior herbaceous formations of a large 
part of Africa and tropical Asia." (Virot, pp. 

The examination of the lists of species reveals, 
in addition, a deficiency in leguminous plants. 

t Presented by R. Heim. 

Papy ( 1954) also points out the absence of prairial 
leguminous plants, endemic species as well as 
those introduced in ancient times, in the group 
known as the Society Islands. 

Thus, what the authors here call "secondary 
savannahs" would be nothing but anthropic 
herbaceous clearings. 

Are there no "primary savannahs" in existence 
in the Pacific islands? Have they, incidentally, 
ever existed ? 

We will try to give here both an analysis of 
this question and a possible interpretation with 
the help of elements of isolated value. 

Two hemicryptophytic graminaceous plants to 
be found to a certain extent everywhere in the 
Pacific Region should have our attention : Themeda 
triandra Forsk and Heteropogon contortus Roem. 
and Sch., two andropogoneous plants. 

Themeda triandra is exclusively paleo-tropical 
in its distribution; Heteropogon contortus is pan- 
tropical, present in the old and the new tropical 

J. Lebrun (1947) proposed the setting-apart of 
a superior group in the phyto-sociologicous 
African hierarchy, the Themedetalia triandrae 
order. This order, of paleo-tropical value, would 
in East Africa include an alliance, the Themedion 
triandrae afro-orientale. (Lebrun). Among the 
vegetable associations included herein, Lebrun 
has studied and defined an association with 
Themeda triandra Forsk. and Heteropogon con- 
tortus^ distinguished as Themedeto-Heteropogone- 
tum, in accordance with the appellative rules of 
nomenclature in phyto-sociologicous subjects. 

The extension of the genus Themeda in the 
whole of the inter-tropical and sub-tropical zone 
of the old world contrasts with its total absence 
in America. 

Very generally speaking, all the species of 
Themeda are orophileous or sub-orophileous, at 
least on approach to the equator. In relatively 
high latitudes, they become "planitiary" (habitat 
in plains). 

The "prairies" of Themeda triandra or of 
T. quadrivahis Kunth characterize a prairial 



pyrophytic type of humid estival tropical climate. 
This type, as we shall see later, seems to be very 
ancient in the world. The discontinuity in the 
area of distribution of two of these species is a 
partial proof of this antiquity. 

The existence of a Themedion triandrae afro- 
orientate does not however presume the existence 
of an exclusively West African alliance, at least 
in the sense of a Themedion triandrae afro-occi- 
dentale. Lebrun (1947) gave an explanation on 
this subject, suggesting for this latter region a 
vicarious order in relation to Themedetalia and 
based on another Andropogon Hyparrhenia 
diplandra Stapf. 

We have shown (R. Porteres 1951) that, if 
traces could be found in West Africa of the 
Themedetalia order, of the Themedion alliance 
and even of a Themedeto-heteropogonetum asso- 
ciation, this has now all collapsed and is rclictual. 
The Hyparrhenietalia partially took its place, 
not as a floristic vicariant, but as an eco-vicariant. 
We have clearly indicated the ecological contrast 
between these two orders and have demonstrated 
the relictual state of the Themedetalia in West 

Taken generally, the prairial formations of 
Th. triandra can be defined as "pyro-climatic" 
and those of Hyparrhenia diplandra as "anthropo- 
pyro-edaphic" (R. Porteres, 1951). 

In East Africa, the floristic individuality of the 
Th. triandra and //. contortus grouping had al- 
ready been recognised by R. Staples (1926) in his 
pastoral experiments on the cultivation of the 
South African Veld. For Staples and Lebrun, 
the grouping tends towards a seasonal contrast 
which results in a marked periodicity of the 
vegetation of this prairial type and thus favours 
its maintenance. The periodical spreading natural 
fires facilitate the extension of the establishment 
of Th. triandra and maintain, in particular, the 
association with //. contortus. 

At Dahomey it is said that a residue of this 
association exists (Porteres, 1951). 

Certain differences in the ecological behaviour 

of the two species could dislocate the association. 
Th. triandra tends towards lighter and better 
porous soils than H. contortus. The latter species 
at present pan-tropical, appears to be more 
aggressive, probably on account of its facilities for 
the scattering of the seeds by man and animals 
(spikelets which catch on with their awns). 
Th. triandra is less aggressive, and this perhaps 
explains its absence in America. 

In certain countries of the Old World, Th. 
triandra is replaced by T. australis (South Africa), 

T. quadrivalvis (Madagascar, part of India). Life 
associated with Heteropogon contortus is also 
to be noted. 

The presence in the Pacific lands, either of 
//. contortus alone, or of Th. Triandra alone, 
or of the two species together, raises in our 
opinion the problem of the existence or the non- 
existence of primeval herbaceous groups, of 
"primary savannahs." 

If these species are of recent introduction, the 
problem does not arise; they would indicate a 
tendency towards the regression of the potential 
vegetation of the "present secondary savannahs" 
which would thus be approaching a pyrophile 

If these species are present since very remote 
ages, this means that pyrophytic prairies existed 
and formed part of the natural landscape of the 
Pacific islands. 

The fact that certain isolated islands or groups 
of islands in the Pacific, such as the Society 
Islands, possess only Heteropogon contortus and 
do not possess Th. triandra give us the impression 
that the pyrophytic conditions of these islands 
are recent and that primary savannahs have never 
existed there. 

However, the establishment of the vegetation 
of all these islands is of recent origin, by the 
covering of uplifts (or volcanic lava stretches) 
with vegetation, dating from the Pliocene age, 
or even the Pleistocene or the Holocene age. 
It is thus impossible to deduce anything much, 
except perhaps that it was man who destroyed 
forest vegetation and who favoured the her- 
baceous clearings which fires were beginning 
to ravage. 

Quite different is the problem of New Caledonia 
where we ascertain the presence, together with 
H. contortus, of Th. triandra. Other species of 
Themeda exist in New Caledonia: Th. ciliata 
which is known to be annual and of recent intro- 
duction; Th. gigantea, an annual para-littoral 
species which does not interest us here and which 
was probably brought in anthropically from India 
over Melanesia. 

Is the Th. triandra in New Caledonia a species 
which is the outcome of natural ancient distri- 
bution? Was it part of the normal surface of 
this island in archaic times ? 

The genus Themeda covers a total paleo-tropical 
area reaching from one extremity of Africa over 
India and Australia and into New Zealand (?). 
Themeda triandra, we may say, is to be found in 
the whole of the area covered by the genus. The 
species includes numerous variations, certain of 



which have been established as independent 
species, and the surviving varieties of which are 
apparently well isolated morphologically; this 
isolation is also, correlatively, of a geographic 
order and reaches as far as the low systematic 
levels, so that it has been possible to use it for 
various interpretations in Africa (Stapf 1919, 
Porteres 1951). 

Incidentally, the genus Themeda (and in parti- 
cular the species Th. triandra in the wide sense) 
particularly merits a very thorough study at 
various levels: systematic, geographic, caryologic, 
cytogenetic, physiological, ecological, sociological, 
and pastoral. 


It is not possible to know whether the Pacific 
lands carried balanced herbaceous formations of 
primeval origin in ancient times. One now finds 
herbaceous clearings which have been introduced 
historically. None the less, the existence of the 
pan-tropical species Ileteropogon contortus Roem 
and of the paleo-tropical species Themeda triandra 
Forsk in these meadow lands permit us to assume 
either a prairial advance towards a pyrophytic 
stage or a relictual state of pyrophytic prairial 

To know whether primary prairies ever existed 
in the Pacific lands is a problem which is far from 
being solved. 

In the case of the Central Pacific islands which 
are geologically young, one should perhaps put 
forward the view that they have never existed; 
but the case of certain outer, geologically ancient, 
chains of islands which belong to the Tertiary Age 
of Indo-mclanesian Asia or of the Australian 
continent, or of an antarctic continent, may be 


Staples, R.R., 1926, Experiments in Veld manage- 
ment, Union S. Afr. Dept. Agric. Sc. 
Bull., 49. 

Lebrun, Jean, 1948, La Vegetation alluviale au 
Sud du lac Edouard, Bruxellcs, Jl: 

Papy, H. Rene, La Vegetation dcs lies de la 
Societe et de Makatea, Vol. I, 386 pp. (in 
Geographic forest, du Monde, Tome V, 
Sect. 2, VoL 1, Art III, 26me Partie). 

Guillaumin, A., 1948, Flora analytique et synop- 
tique de la Nouvelle-Cale"donie-Phanero- 
games. Vol. I, 369 pp., Paris. 

Guillaumin, A., et Virot R., Contributions a la 
Flora de la Nouvelle-Caledonie. CII 
Plants recoltees par M.R. Virot. Mem. 
du Museum National d^Histoire Naturelle. 





Ciha* Basle, Switzerland. 

Attempts to describe thoroughly and classify 
the New Caledonian vegetation were started only 
about 30 years ago. Last year, an important 
study by R. Virot was published, giving for the 
first time a synopsis of the vegetation groups of 
the whole island. Together with the studies by 
Daeniker, Sarlin and other authors, we now 
have substantial data regarding the composition 
of several groups of plant species. However, 
precise observations of documentary nature are 
partly lacking with regards to frequency, arrange- 
ment and development of species within the bio- 
coenoses. The works of Dansercau, Richards 
and Schmid set out the lines along which such 
studies should be conducted, which are of funda- 
mental importance to the knowledge of the ecolo- 
gy of the species and its application in forestry, 
agriculture and livestock. 

By adopting the method described by Schmid, 
we endeavoured to get an idea of the structure of 
parts of some biococnoses. Unfortunately, due 
to limited time, all generalized study was impos- 
sible. On another occasion, we described this 
method, making certain changes that appeared 
useful, for the determination of the structural 
features and their illustration. 

The purpose of this paper is to give an idea of 
the structural facts which are recognizable through 
these biocoenological surveys. 

Sarlin mentions the classification of the New 
Caledonian vegetation into 7 distinct formations 
established by previous authors: 

1. Coastal zone with Oceanian flora; 

2. from to 3-400 m the Niaouli Savannah 
( Melaleuca leucadendron) ; 

3. from 400 to 1,000m medium altitude forest; 

4. from 1,000 to 1,500m dry coniferous forest; 

5. the forest galleries; 

6. lower serpentine scrub; 

7. summit scrub. 

Those who know New Caledonia well realize 
that this classification is superficial, incomplete 
and even erroneous (for example the definition 
of the formation under item 4 as "dry forest." 

Virot has chosen the following system to give 
a comprehensive view: 

1. Halophilous formations, including: 

a) mangrove, 

b) herbaceous vegetation of beaches, 

c) belt of so-called "Seaside" trees; 

2. Non-halophilous formations, divided into: 
A forest series, including: 

a) coastal climatic forests (coastal hillside 

b) hydrophilous forests of the river banks, 

c) photophilous forests of the marshes, 

d) mesophilous forests of the valleys, 

e) shade-tolerant forests (insufficiently speci- 

f ) oro-nephepiphilous forests, 

g) photo-xerophilous forests; 
A series of shrubs, including: 

h) edaphic sclerophilous-xerophilous scrub, 
i) orophilous scrub; 
A herbaceous series, including only: 
k) paludal groups. 

Of course, this classification will only be provi- 
sional since it does not take into account the mixed 
floristic features which constitute the vegetation 
of different regions of the island. Despite this 
defect, Virot's classification shows considerable 
progress when compared with previous classifi- 

Our structural studies were carried out mostly 
in the formations which Virot calls vallicol-meso- 
philous and oro-nepheliphilous, which are 
obviously very mixed formations. By way of 
comparison, we will also discuss a survey made in 
a secondary coastal forest. On the other hand 
we will not talk about shrub vegetation of ser- 
pentine areas which we have not been able to 
study in detail. 

The forests in which we were most interested 
because of their aspect are those with a predom- 
inance of Neo-caledonean Fagaceae, of Trisyn- 
gyne spp. They are distinct from other types of 
forests by the fact that only one species forms, 
nearly exclusively, the dominant tree stratum. 

Our surveys Nos. 6, 9 and 10 were made in 
Trisyngyne codonandra Baill. forests; survey No. 
7 in a 7>. Balansae Baill. forest. All the fragments 
studied are located on peridotitic mountains 
quite near to one another: the farthest apart are 
separated by less than 30 km. In spite of this, 
relatively important differences are apparent, 
even between the three Tr. codonandra biocoe- 

Due to lack of time, it will not be possible to 
reproduce here the complete lists of species. 



Their study shows that the floristic composition 
is very variable in Tr. codonandra forests. Among 
all the species found in the 3 surveys, there are 
only two species, Tr. codonandra itself and 
Symplocos cf. Pancheri, found in all three surveys. 
Five species are common to surveys 6 and 9, 
eight to surveys 9 and 10, while surveys 6 and 10 
have no particular species in common. 

It is interesting to see that survey No. 7, in 
spite of the dominance of another species of 
Trisyngyne, seems to connect the other fragments 
of vegetation. In fact, surveys No. 7, 9 and 10 

have 3 species in common, and surveys 6, 7 and 
9 two species. The total of the plants approxi- 
mately identified in the 4 fragments studied is 
about 150 phanerogamic species. The coincidences 
found are therefore rather poor, and it will be 
admitted that the floristic-statistics method is 
not of very great use in these circumstances. 

Now let us proceed to the information received 
on biocoenological structure. By grouping the 
plants together, forming provisional ecological 
types, the following frequency of the different 
types in the biocoenoses fragments occurs : 

Table 1. 

Frequency of provisional ecological types in the vegetation according to the surveys 
(in per cent of small squares of the surveys): 



8 = 

k = 










trees more than 1 2 m high (number) 

shade produced per crown 

trees more than 8 to 12 m high (number) 

shade produced per crown 

trees less than 8 m high 

trees with tufts of leaves, 2 to 8 m high, 

few branchings 

trees with tufts of leaves, less than 2 m high, 

few branchings 

prostrate trees 

shrubs more than 2 m high 

shrubs less than 2 m high 

woody lianes or lianes with stiff stems 

prostrate shrubs 

erect herbs 

herbs forming dense clumps 

herbs forming rosettes 

herbs forming stems 

epiphytic herbs 

bryophytes on the ground 

bryophytes on wood (dead or living) 


saprotic mushrooms 


no data 

no data 


























































To these figures must be added those of young plants which, according to their state of growth, parti- 
cipate in the composition of the various biocoenoses strata: 

Young plants more than 1 m high 
Young plants, 20 -100 cm high 
Young plants less than 20 cm high 







The most noticeable differences are found in the density of class c trees (including class e and 
the young trees over 1 meter high) and in that of non-epiphytic undergrowth (including young plants 


of less than 20 cm high), 
are as follows: 

Class c etc. 
Class p etc. 


The total percentage of participation of these 2 categories in the 4 surveys 








Therefore, surveys Nos. 7 and 9 present a much more developed undergrowth than Nos. 6 and 10. 
This fact is also a result of the number of small squares entirely without any vascular plants (in % of 
the total of squares): 

6 9 10 7 




These differences are due to the growth of the 
dominant tree strata, producing more or less 
intense shade according to the density of the 

The frequency of the shrubs follows the same 
lines, but their absolute number is very small, 
so that they only play a minor part in the structure 
of the vegetation as a whole. 

The proportion of lianes is almost similar in 
the 3 surveys Nos. 6, 9 and 10, but there is an 
abundant presence of young creepers (Frcycinctia 
sp., Smilax sp. and Alyxia sp .) in survey No. 6. 

Jn the participation of young plants, it is ad- 
visable to compare the number of adult and young 
subjects of various species. These observations 
give valuable information on the probable 
development of the biocoenoses in question. The 
lack of young plants of dominant species can 
induce a change in the structure of the whole 
vegetation, whilst an abundance of young plants 
is a certain guarantee for the maintenance of 
population. Below are the most striking examples 
of both statements, found in our surveys: 

a) Very abundant young plants, compared to adult 

young adult 

Survey No. 6 

Trisyngyne codonandra \ 39 3 

Tree No. 40 22 1 

Survey No. 9 

Araucaria muelleri 108 

Palm tree No. 24 311 

Survey No. 10 

Trisyngyne codonandra 46 3 

Rubiaceae No. 29 23 1 

Survey No. 7 

Trisyngyne balansae 652 8 42 10 

Guttiferae No. 32 33 

b) Abundant adult plants, few young plants: 

young adult 

Survey No. 9 

Trisyngyne codonandra 4 7 

Cunoniaceae No. 9 2 14 

Myrtaceae No. 3019 3 17 

Survey No. JO 

Sapindaceae No. 34 08 

Survev No. 7 

Hibbertia sp. No. 14 8 

Montrouziera sp. No. 4 117 

Leucopogon dammarifolius 6 14 

In surveys Nos. 6, 10 and 7, seed reproduction 
of the dominant species is well favoured. On 
the other hand, its future is uncertain in survey 
No. 9. Here, the young Araucaria muelleri plants 
occupy a very important place in the vegetation 
of the lower strata although there are no adult 
trees of this species in the square of the survey or 
in its immediate neighbourhood. 

However, the abundant seed production and 
germination are still not sufficient to guarantee 
the stability of the stock of plants. Furthermore, 
development must also take place without any 
particular hindrance, whilst allowing for a succes- 
sive elimination of young plants by competition. 
In surveys Nos. 6 and 10, this phenomenon is 
apparently present since there are young plants 
of all sizes. The case of survey No. 7 is different : 
beside 8 adult trees, we find 543 young plants of 
10 cm or less, 105 of 11-20 cm, only 4 plants 
between 21-30 cm and no taller plants. Beside 
an irregular seed production from year to year 
which seems likely, it is probable that further 
growth of the young plants is hindered when they 
have reached a certain age, although we cannot 
indicate the factors responsible for this. 

We should like to mention here a study made by 
Mr. Letouzey, Conservateur des Eaux et Forets 



in French Camerons, on the Lophira alata forest 
of the coastal zone of the Camerons. There, 
the Lophira alata has the same symptoms of com- 
petitive insufficiency in its youth, and the author 
presumes that without protected artificial regen- 
eration, the forest composition would change. 

The interpretation of survey No. 9 is rather 
delicate. The four young plants of the dominant 
species are 20, 30, 40 and 40 cm high respectively. 
An adult plant of 6 meters only has not yet 
attained its optimal growth. However, the massive 
growth of young Araucarias between 20 to 160 
cm high seems to show that this forest will be 
changed as and when the old Trisyngync die. This 
transformation is apparently favoured by a rather 
weak shade produced by the adult Trisyngync, as 
their crowns are not contiguous, and by the fact 
that other species participate in the formation of 
the dominant stratum (A gat his ovata, Salauopris 
sparsiflora and Rhodamnia andromcdoides). In 
fact, at this high altitude, Trisyngyne does not 
seem to be at its best, and its position seems 
to be endangered by the strong competition of 
other plants. Changes of secondary order in 
ecological conditions may consequently easily 
provoke structural changes and accelerate the 
dynamic tendencies of the biocoenose. 

One last structural component on which we 
would like to draw attention is the distribution 
of the species in the area surveyed. Beside 
plants like Phclline lucida and Myodocarpus 
fraxinifolius of survey No. 6, or Leucopogon dam- 
marifolius of survey No. 7, which are all quite 
regularly distributed over the whole area, there 
are others such as Rubiaceae No. 29 of survey 
No. 10, or Lophoschocnus montisfontium of sur- 
vey No. 9, the former located in one corner only, 
the latter separated in three distinct groups. 

These irregularities can be caused by two com- 
pletely independent factors: 1. the character- 
istics of the biological system of dispersion of 
fruits or seeds, as well as the possibilities of a 
vegetative reproduction, or, 2. conditions of the 
biocoenological medium allowing the develop- 
ment of plants in one area, and hindering it in 

The first factor is apparently responsible for the 
grouping of the plants of Rubiaceae No. 29 men- 
tioned: all the young plants are grouped around 
an adult tree. To a certain extent, tree No. 40 
of survey No. 6 and Spiraeanthemum sp. No. 
2 of survey 10 show similar phenomena. On the 
contrary, with regards to Lophoschoenus montis- 
fontium, their relation to the medium seems more 
important: it is apparent that it is found mostly 


under the spaces left open by the crowns of domi- 
nant trees. The contrary could be said regarding 
Phelline lucida of the same survey (No. 9) which 
seems to follow the shade of the trees. Therefore, 
this species shows that the group forming tenden- 
cy is subject to the influence of the vegetation 
itself, having a homogenous distribution in a 
forest of regular shade, forming groups in a 
forest of irregular shade. 

The presence of groups of certain species, in 
relation to other plants, is very interesting for 
the classification of vegetations. However, care 
should be exercised in order to avoid overesti- 
mation of the value of the groups. Daeniker has 
always insisted on the necessity of considering 
groups within a wider framework, to look at them 
as fragments of a mosaic. Temporary dominant 
or subdominant conditions caused or suffered 
during certain states of individual growth of a 
species can be very important, without justifying 
however a classification of biocoenoses into 
smaller defined units. In this respect, we recall 
the lively discussions which took place during 
the last International Congress of Botany in 
Paris concerning the existence of tropical asso- 

Taking our structural analysis into account, 
we will not hesitate to put surveys 6 and 10 
together as having both the same type of biocoe- 
noses, despite the absence of common species in 
the under-growth and herbaceous stratum. This 
type is characterized by one dominant stratum of 
trees of equal height, reaching 12-15 m. The 
lower strata are not very developed. There 
seem to be minor differences only in the compo- 
sition of the stratum of small trees (presence of 
many trees of provisional ecological type e) as 
well as in the frequency of young plants. 

Survey No. 9 is clearly different in its structure. 
Dominant adult trees do not reach the dimensions 
of the Trisyngyne of the two afore-mentioned 
surveys. The crowns do not form a contiguous 
cover and consequently, the undergrowth and the 
herbaceous stratum are much more developed. 
This structure is emphasized by the abundance of 
young plants. Analysis of the fragments of 
neighbouring vegetations will doubtless show 
that this is a mosaic stone forming a transition to 
distinct biocoenoses of higher altitudes. 

Survey No. 7 belongs to a third type of biocoe- 
noses. The dominant trees reach 12-15 m in 
height, but without forming a complete cover at 
crown level. The foliage of TV. balansae produces 
less shade than that of Tr. codonandra. Therefore 
the undergrowth and the herbaceous stratum are 



well represented and heliophilous shrubs such as 
Leucopogon, Dracophyllum and Hibbertia, as 
well as the heliophilous orchid Eriaxis rigida, 
can be found. Contrarily to what we find in 
survey No. 9, there is a lot of herbs with a tendency 
to form aerial stems without leaves (Lophos- 
choenus spj, while rosette herbs are lacking. 

The common characteristics existing between 
the four surveys cannot fail to be noticed: the 
dominant stratum reaching a certain height above 
the ground and formed essentially by one species 
alone, with, at most, 2 to 4 species reaching the 
same level, but isolated between the Trisyngyne. 
The most frequent families by number of individ- 
uals are the following: Fagaceae, Polypodiaceae, 
Orchideae, Cyperaceae, Sapindaceae, Rutaceae, 
Aquifoliaceae and Guttiferae (thus, three families 
comprising mostly herbs). 

Now let us compare briefly another survey 
with the four fragments discussed: No. 8, situated 
near No. 7, on a steep slope of a pcridotite moun- 
tain. Even at a superficial glance a complete 
difference from the afore-mentioned biocoenoscs 
is apparent. 

Above a tree stratum forming a continuous 
dome but unequal as to its level above the ground 
(between 10-15 m in general) a few isolated trees 
grown up to a height of 20-40 m (Agathis lanceo- 
lata, Araucaria hernicri, Alhizza sp. and other 
species). Their ecological influence is unim- 
portant compared to the trees forming the dome. 
Despite the dense shade produced by the latter, 
the stratum of trees of less than 8 m high (types c 
and e) is represented in abundance, and small 
shrubs (type k) are more frequent than in any 
Trisyngyne forest surveys. Herbs are also found 
in greater abundance than in fragments of shaded 
biocoenose of Tr. codonandra Nos. 6 and 10. 
Finally the very high number of young plants 
(33 % of the square are young plants higher than 
1 m, 91% young plants of 20 to 100 cm, 77% 
young plants smaller than 20 cm) indicates that 
this biocoenose is well balanced and that its 
participants are well adapted to the given condi- 

This is explained by the very different floristic 
composition of this forest, compared to previous 
surveys. The most represented families by number 
of individuals are in this case: Rubiaceae, Pan- 
danaceae, Arabiaceae, Guttiferae, Euphobiaceae, 
Myrtaceae, Laureaceae and Myrsinaceae, i.e., 
families of trees found in all dense tropical forests 
of the world. The undergrowth includes a rather 
large number of sciaphilous species, specially 
a Phyllanthus and two Psychotria. The Freyci- 

netta genus is also well represented, specially 
among young plants. Amongst the genera en- 
countered in the Trisyngyne biocoenoses, only 
Phelline and Myodocarpus fraxinifolius are found 
in greater number, hence, trees which are also 
found under shade in survey No. 6. 

Finally, on 100m 2 , we have found 14-16 species 
in the strata of trees of over 8 m high, and it is 
certain that this number would increase if the area 
observed was extended. 

Therefore the structure of the biocoenose frag- 
ment of survey No. 8 is that of a tropical forest. 
By its complexity and the great amount of species 
it contains, it is clearly distinct from Trisyngyne 
forests. Of course, it does not correspond to 
dense forests of the type of equatorial rain forest 
of the big continents, but it is noticeable that the 
general idea, if I may call it so, is similar. Besides, 
we have seen very similar forests from the struc- 
tural point of view in Central America, in East 
Cuba and on the Eastern slope of the Drakens- 
berg, South Africa. 

Trisyngyne forests, with their simpler structure 
and predominance of one species only in the 
higher tree strata, are more reminiscent of forests 
of temperate regions. However, it would be 
unwise to use this term in defining any vegetation 
of New Caledonia. In fact, the geological evolu- 
tion of this island, the phylogeny and epi-ontology 
of its vegetation are so different from what is found 
in temperate countries that we would prefer to 
create a new expression: "paratropical forest" 
next to tropical vegetation properly speaking. 

To add another type of structure, we would 
like to mention briefly the study of a fragment 
of a secondary forest, greatly influenced by man, 
located on the slopes of Ouen Toro near Noumea. 
The ground is of sedimentary origin, and the 
vegetation varies from very impoverished grass- 
lands to regenerated forests, as described by 
Sarlin in his work. 

. In the fragment studied, we again find a domi- 
nant tree stratum, only reaching 8 m in general, 
composed nearly exclusively of Acacia spirorbis. 
A few trees of other species are mixed therein: 
Casuarina cunninghamiana and Melaleuca leuc 
adendron. Under the shade of these trees, 
the crowns of which are close enough together 
without producing however a too intense shade, 
-the small trees and shrubs are few. On the con- 
trary, the herbaceous stratum is very developed 
(specially Bidens pilosus, Plectranthus parviflorus, 
Rynchelytrum roseum and other Graminaceae and 
Composites),and the ground is abundantly covered 
by young Passiflora suberosa plants, Observing 



the distribution of herbs throughout the survey, 
it is apparent that the structure is not firmly 
established, because the herbs are generally lim- 
ited into groups, with small interference of species. 
Therefore, we are dealing with initial stages rather 
than with a normal vegetation competition. 

As I said in the beginning, this summary of our 
biocoenological studies of New Caledonia is not 
sufficiently complete to give you a general view of 
the types of structures found on this island. We 
have spoken neither of swamp forests on sedi- 
mentary grounds which are distinct from those 
found on peridotitic grounds, nor of the Niaouli 
savannah (Melaleuca leucadendron ) , which has 

its own many difficult problems as to its original 
genesis, problems which perhaps only a detailed 
analysis of the structure could solve. We have 
not discussed the following points: the shrubby 
"bush" and the serpentine vegetations, with their 
various strata and types; to what extent are they 
natural, are they caused by fire or by cutting? 
We were unable to compare the vegetation of 
the peridotitic, schistose and gneissic mountains, 
and we have not established the characteristics 
of various altitude zones. All this would require 
much more time for field work than we had, and 
the only thing we seek to accomplish by this paper 
is to show the Wciy in which one can attempt to 
solve these questions. 






The Pennsylvania State University, University Park, Pennsylvania, U.S.A. 

Coastal forest types in western Washington and 
Oregon resembling tropical forests by their 
luxuriant epiphytic growth on trees, have been 
called "rain forests." Generally these rain forest 
types are confined to the Fog Belt Subregion. 


The unique physiography of the Fog Belt Sub- 
region consists of high mountain ranges directly 
parallel to the Pacific Coast blocking the path 
for southwesterly ocean winds. These mountain 
masses (Coast Range and the Olympics) form a 
barrier of 3,000 to 8,000 feet elevation forcing 
these moisture-laden storms to rise and condense, 
resulting in the highest orographic precipitation 
for the whole North American continent. Several 
large rivers have cut through the Coast Range 
creating wide valleys open to these ocean winds. 
Glaciers advancing down from the Olympics have 
scoured other deep, steep-sided, U-shapped valleys, 
in places sometimes more than a mile wide. 

These glacial valleys are essentially level with 
a gradient of about 1 %. The highest precipita- 
tion occurs at the end of the 2-30 mile long and 
broad valleys around 1,500-2,000 feet elevation, 
sometimes resulting in an annual precipitation of 
more than 300 inches. 


The proximity of 2-20 airline miles to the 
Pacific Ocean creates an ideal marine climate in 
this narrow Fog Belt Subregion, extending some 
500 miles along the Pacific Coast, from California 
into Alaska. 

General characteristics of this marine climate 

(1) A very narrow fluctuation of temperature 
extremes hardly exceeding 20F annually with 
an average annual temperature of 49 - 50F. 

(2) The longest recorded frost free period 
extending well over 200 days, ranging from 180- 
310 days which is exceptional for the 44-49 paral- 
lel northern latitude. 

(3) Consequently, it has no snowfall directly 
near the ocean. Six miles further inland in the 

valley bottoms snow depth increases from a few 
inches to two feet and at a distance of 15 
miles inland from 2 to 4 feet. Above 1,000 feet 
elevation, annual precipitation in the form of 
snow can even amount to 300 inches at high 

(4) No appreciable drought period in the 
summer. During the early morning hours incom- 
ing ocean fog will create considerable fog drip 
within the rain forest preventing these forest types 
to dry out. Forest fires which commonly ravage 
more inland forest types have never been able to 
penetrate into these rain forest types. 


Rain forest types cover river flats and the foot 
hills up to about, 1,000 feet elevation constituting 
of two, possibly more, alliances of coniferous 
forest types dominated by the Sitka spruce and 
western hemlock, and of deciduous forest types 
dominated by the big -leaf maple and red alder. 

The American rain forest types are character- 
ized by: 

(1) Unusual tree heights and sizes. The largest 
and highest individuals of Douglas fir, Sitka spruce, 
red cedar, and hemlock are all found within the 
rain forest. Tree heights of 300 feet and more are 
not uncommon in virgin stands of sheltered 
valleys with tree ages of about 2,500 years. 

(2) Buttressed tree bases and collonades. 
Sitka spruce and hemlock regeneration usually 
starts upon rotting logs and stumps. Seedling 
growth is very slow on these substates until its 
roots strike mineral soil. Roots become heavier 
and fuse in order to support the increasing tree 
size. Gradually, the nurse-log decomposes, leaving 
a collonade of trees with buttressed bases and 
exposed root collars. 

(3) Luxuriant epiphytic growth on stems and 
branches of trees. In temperate forests, epiphytic 
vegetation is rarely encountered. Low tempera- 
ture and occasional dessication in the summer 
have limited the epiphytic flora to a few crypto- 
gams, lichens, bryophytes, and pteridophytes. 
No profusion of vascular epiphytic plants, so 

t Presented by F.E. Egler. 



typical of tropical rain forest, exist. Trees may 
become sometimes so heavily laden with an 
upholstering of rain-soaked bryophytes that they 
are susceptible for windthrow. 

The coniferous forest alliances can be divided 
into four main forest types: 

The Sitka Spruce - Hemlock forest type. This 
is the most common rain forest type covering 
river flats, slopes, and ridges. Its tree layer is 
dominated by Picea sitcheniss and Tsuga hetero- 
phylla usually in an intensive mixture, however 
locally pure stands of one of these dominant trees 
are also found. Its shrub layer is dominated by 
Acer circinatum or Rubus spectabilis particularly 

in river flats and on lower slopes. Its herb layer 
is characterized by the dominance of Polystichum 
munitum, Oxalis oregana, and occasionally 
Tiarella trifoliata. 

The characteristic species composition includes: 

the group of character species of this forest 

Arceuthobium tsugen- Moneses uniflora 


C V/V? ton la un (flora 
Cornus canadensis 
Disporum smith! i 
Mcnziesia ferruginea 

Picea sitchensis 
Rubus pedatus 
Tsuga heterophylla 
Vaccinium alaskaense 
Vaccinium ovalifolium 

Particularly the stands on river flats, lower and 
middle slopes are characterized by the common 
moisture indicators: 

Acer circinatum 
Athyrium felix-femina 
Blechum spicant 
Disporum oreganum 
Dryopteris dilatata 
Fesfuca subulifiora 
Luzula parviflora 

Melica subulata 
Montia sibirica 
Oxalis oregana 
Polystichum munitum 
Rubus spectabilis 
Sambucus callicarpa 
Stachvs cilia ta 
Tiarella trifoliata 

Stands on upper slopes are generally character- 
ized by the lack of moisture indicators and by the 
abundance of: 

Gaultheria shallon Linnaea borealis 

The Coastal Sitka Spruce forest type. This 

forest type is limited to bluffs and west slopes 
directly facing the Pacific Ocean. Dominant 
trees of Sitka spruce, hemlock, and red alder 
are stunted by wind. Its shrub layer is usually 
dominated by Gualtheria shallon, sometimes 


on the lower slopes by Rubus spectabilis. 
The characteristic species composition consists 


Alnus ruhra 
Arceuthobium tsugense 
Lonicera involucrata 
Picea sitchensis 

Thuja plicata 
Tsuga heterophylla 
Vaccinium ova turn 

together with most of the common moisture 

The Sitka Spruce -Lodgepole Pine forest type. 
This forest type is confined to areas with sandy 
soils and dune formations in the vicinity. 

Its characteristic species composition consists 


Gaultheria shallon 
Lonicera involucrata 
Mains rivularis 
Picea sitchensis 
Pinus contorta 

Thuja plicata 
Tsuga heterophylla 
Umbel la ria calif or nica 
Vaccinium ova turn 

with hardly any of the common moisture 

The Sitka Spruce Swamp forest type. Meander- 
ing rivers create extensive swamp forest areas in 
their deltas. Trees are situated on humps within 
the swamp. Its tree layer is rather open, charac- 
terized by the dominance of Sitka spruce and red 
alder mixed with an occasional hemlock; its 
shrub layer is dominated by Rubus spectabilis. 

Its characteristic species composition consists of: 

Alnus rubra 
Cardamine angulata 
Car ex obnupta 
Chrysoplenium glecho- 

Lonicera involucrata 
Lysichitum camtschat- 

Oenanthe sarmentosa 

together with the character species of the 
Sitka Spruce - Hemlock forest type: 

Menziesia ferruginea 
Moneses uniflora 

Vaccinium alaskaense 
Vaccinium ovalifolium 

with the common moisture indicators: 

Athyrium felix-femina 
Blechnum spicant 
Dryopteris dilatata 
Luzula parviflora 

Polystichum munitum 
Rubus spectabilis 
Sambucus callicarpa 
Stachys ciliata 
Tiarella trifoliata 


In the deciduous rain forests, the Riverflat 
Maple and the Red Alder forest types are disting- 
uished. The Riverflat Maple forest type is noted 
for its luxuriant epiphytic cover, being considered 
usually the veritable rain forest. In virgin stands 
the dominant bigleaf maple is mixed with Sitka 
spruce, red alder, cottonwood, hemlock, and 
grand fir, but in second growth stands usually 
only the maple is present. A luxuriant cover of 
bryophytes and vascular plants cover the soil, 
rotting logs and tree trunks while a dense herb and 
shrub layer will add to its jungle-like appearance. 

However, on the Olympic Peninsula these 
Riverflat Maple forests have a rather open and 
park-like character, lacking chiefly the dense layer 
of high herbs and shrubs. The reason for this is 
the presence of some 4,000 Roosevelt elk (Ccrvus 
canadensisvar: roosevelti) wintering in these rain- 
forest valleys inside the Olympic National Park. 
As the Roosevelt elk is gregarious, frequently 
banding in herds from 20 to 100 animals, the 
effects of elk herds' browsing and trampling are 
significant. The lack of high herbs and shrubs and 
the introduction and abundance of grasses and 
grazing indicators have given this forest type an 
overbrowsed appearance. Since 1935, elk cxclo- 
sure plots have been established within the 
Olympic National Park. At the present the vege- 
tation inside these fenced plots is characterized 
by a tall herb layer and a dense shrub layer of 
Acer circinatum, Sambucus callicarpa^ and 
Rubus spectabilis indicating its natural appear- 
ance without the overbrowsing effects of the 
Roosevelt elk. In other parts of the Fog Belt 
Subregion where no Roosevelt elks are present, 
deer will frequently graze in the Riverflat Maple 
and Red Alder forest types, so that grazing 
indicators and grasses form a characteristic com- 
ponent of these deciduous rain forest types. 

The Riverflat Maple forest type. The charac- 
teristic species composition of this forest type is 
comprised of: 

in its tree layer: 

Pseudotsuga taxi folia 
Selaginella oregana 

Thuja plicata 
Tsuga heterophylla 

Abies grandis 
Acer macro phyllum 

A Inns rubra 
Picea sitchensis 
Populus trichocapra 

in its shrub layer : 

Acer circinatum (dom.) Holodiscus discolor 
Corylus californica Oplopanax horridus 


Samhucus callicarpa 
Tax us brevi folia 

Osmaronia ccrasiforis 
Rubus spectabilis 

in its herb layer: 

the group of common moisture indicators: 

Acer circinatum 
A thyrium feliz-femina 
Blechnum s pi cant 
Dent aria tenella 
Disporum oreganum 
Dryopteris dilatata 
Festuca subuliflora 
Luzula par vi flora 

Melica subulata 
Montia sibirica 
Oxaiis oregana 
Rubus spectabilis 
Sambucus callicarpa 
Stachys ciliata 
Tierella trifoliata 
Trientalis latifolia 

the group of riverflat moisture indicators: 

Asarum caiulatum 
Circaea pad fie a 
Dicentra Jormosa 
H vdrophyllum tenuipes 
Mitella caulescens 
Nemophila par vi flora 
Rosa nut k ana 



Tell in HI grandiflora 
Tolmiea menziesii 
Urlica Ivallii 
Viola glabclla 

the group of grazing indicators: 

Adenocaulon hi color Calium tri/idum 

Agrostisstoloniferajexarala Osmorhiza nuda 
Bromus sitchensisjvulgaris Poa kelloggii 
Cardamine oligosperma 
Carex leptopoda 
Deschampsia elongate/ 
Galium aparine 

the group of Riverflat Maple forest character 

Ranunculus bong- 


Stellaria crispa 
Trisetum cernuum 

Carex hendersoni 
Car ex mertensii 

Elymus glaucus 
Galium oreganum 

Epiphytes on trees are comprised of a large 
group of bryophytes and lichens of which the 
most common ones are : 

Antitrichia curtipendula 
Camptothecium lutescens 
Claopodium crispijolium 
Eurhynchium oreganum 
Frullania nisquallensis 
Homalothecium nuttallii 
Hylocomium proliferum 
Hypnum subimponens 
Lobaria oregona 
Metzgeria conjugata 
Metzgeria pubescens 
Mnium menziesii 

Mnium punctatum 
Mnium venustum 
Neckera douglasii 
Neckera menziesii 
Pore I la navicularis 
Pseudoiso thecium 






A few plants are also characteristic epiphytes: 
Montia heterophylla Selaginella oregana 
Polypodium vulgare 

The Red Alder forest type. It occupies similar 
habitats as the former forest type and is usually 
in a successional transition towards it. Charac- 
teristic for this forest type is a dense, even-aged 
stand of Alnus rubra, the almost complete lack 
of any shrub layer in its typical form and the lack 
of a dense epiphytic upholstering of the tree 

trunks. Mostly, the red alder is densely covered 
by crustaceous and foliosc lichens with some 
bryophytes, comprised of: 

the group of common epiphytes: 

Antitrichia curtipendula 
Cetraria glauca 
Frullania nisquallensis 
Ne( '/< cm do u% las ii 
Parmefia enteromorpha 

Pannelia physodes 
Pcrtusaria multipuncta 
P or el la navicularis 
Pseudoiso thecium stol- 

and the group of epiphytes typical for red alder 

Cetraria scutata 
Dimino\veL\ia cirrhata 
Ever n ia prunastri 
Graphis scripta 
Lecanora subfusca 
Ochrolechia tartarea 
Orthotrichum consi- 

Orthotrichum Ivellii 
Parmclia pertusa 
Par me I ia sax at His 
Radula bolanderi 
Radula complanata 

The characteristic species composition of this 
Red Alder forest type is comprised of: 

the group common moisture indicators: 

Acer circinatum 
A thy Hum felix-femina 
Dryopteris dilatata 
Luzula par vi ft or a 
Oxalls oregana 
Polvstichum munitum 

Rhamnus purshiana 
Ruhus spectabilis 
Samhucus callicarpa 
Stachys ciliata 
TiareUa trifoliata 

the group of riverflat moisture indicators: 

Circaea pacifica Osmaronia cerasifor- 

Dicentra formosa mis 

Hydrophyllwn tenttipes To/mica menziesii 

Mitella caulescens Viola glabella 

the group of swamp indicators: 
Cardamine angulata Lysichitum camtschat- 
Chrysoplenium glech- cense 

omaefolium Oenanthe sarmentosa 

Corydalis scouleri 

the group of grazing indicators with the follow- 
ing additional grazing indicators typical for this 
forest type: 

Cirsium lanceolatum 
Equisetum arvense 
Rumex acetosella 

Rum ex obtusifolia 
Stellaria graminea 

and the group of character species of the Red 
Alder forest type: 

Epilohium adenocaulon Pleuropogon refractus 

Glycerin elata 

Mimulus guttatus 

Mite (la ova Us 
Pctasites spcciosus 

Ribes petiolare 
Senecio triangularis 
Yaleriana sitchensis 

The deciduous rain forest types are strictly 
confined to valley bottoms and coves along 
streamlets below 1,000 feet elevation. They are not 
completely restricted to the Fog Belt Subregion 
but may be found locally in the foot hills of the 
Cascades where the annual precipitation is 
unusually high. Both forest types occur on 
alluvial deposits close to the river. The open 
river bars are first colonized by red alder and 

Successional trends of the Red Alder forest type 
towards the climax Riverflat Maple forest type can 
be frequently encountered, but no observations 
have led to the belief that eventually the deciduous 
Riverflat Maple forest will evolve into a conifer- 
ous rain forest type. When the river alters its 
course, it may disrupt forest vegetation and occa- 
sionally Douglas fir may get established in pure 
even-aged stand patches. The forest communities 
in such Douglas fir stands are closely allied to the 
Sword Fern-Douglas Fir alliance. As fire is not 
the tool for opening up the forest canopy for 
future Douglas fir regeneration, only clearing of 
forested land by river erosion and redeposition 
of new land will perpetuate the Douglas fir forest 
community. Otherwise, it will evolve into the 
climax of the Riverflat Maple forest type. 






Hokkaido University, Sapporo, Japan. 

The present project is a research into the forest 
vegetation of Japan. The writer wished to obtain 
a phytosociological description of the various 
forest vegetations, once the tremendous treasure 
of Japan. The primeval forests in Japan have 
been largely destroyed, and this destruction will 
continue until public pressure halts the abuse. 
The writer selected carefully the experimental 
plots represented by the fine forests in the present 
Japan where the excellent physiognomies of the 
natural forest have been fortunately preserved. 

To analyse the construction of the forest 
vegetation, the writer used "the sociation" as the 
synecological unit. He treated it as the species 
in the systematic botany. This idea results from 
the field surveys of the complicated forest veget- 
ation of Japan in these twenty-five years. For 
the description of the sociation, the bisect method 
accompanied by the map, the sketch, the photo- 
graph, and the analytical table was used. It is 
the shortest and the wisest method to express the 
composition of the forest vegetation in the natural 
stand. The writer has attempted to describe 
the vegetation accurately and to analyse precisely 
and carefully the construction of the forest layers. 

Generally speaking, Japan ranges from the 
zone of southern temperate forest to that of the 
northern temperate, each represented by the 
evergreen broad-leaved forest and the summer- 
green one. The former includes the subtropical 
rain forests in its southern points and the latter 
sometimes accompanied by the subarctic forests 
in the northern part. The latter in Hokkaido 
except the southwestern part shows the inter- 
mediate feature between the temperate and the 
subarctic forest, expressing the stronger influence 
of the temperate character. 

From the viewpoint of the vertical vegetation, 
the pure thicket of the Siberian dwarf pine is well 
developed in the alpine belt of the mountain 
ranges in the north from Central HonshQ (the 
main island) through the Be tula Ermani forest. 
The subarctic or needle-leaved forest represented 
by Abies and Picea follows the Ermarfs birch 
forest or directly contact the dwarf pine thickets. 
But it is very characteristic to render the forest 

t Presented by F.E. Egler. 

vegetation in the Japan Sea side by the absence 
of the Picea- Abies forest. 

The climax forests are as follows in the different 
forest zones: 

Subtropical rain 
Salt marsh 

Warm temperate 

Sandy shore 
Coastal district 
Mountain dis- 

Ficus Wightiana 
Kandela Camlet 
Livistona subglobosa 

Pinus Thunbergii 
Machilus Thunbergii 

Abies firma-Tsuga Sieboldii 

Cold temperate forest 

Seashore Quercus dentata 

Light soil Quercus crispula 

Valley Pterocarya rhoifolia-Aescu- 

lus twbina- Cercidiphyllum 



Picea- Abies 

Picea jezoensis- Abies sach- 

Bog Picea Glchni 

Honshu Picea je 20 ens is va r . hondoen- 

s is- Abies Veitchii 
Northern mixed forest 

Sandy shore Quercus dentata 
Plain Ulmus propinqua-A cer mono 

S wany A Inus japonica 

Hill Acer nwno-Tilia japonica 

Valley Cercidiphyllum japonicum- 

Fraxinus mandshurica-Jug- 
lans ailanthifolia 

Up to the present, the following three volumes 
(Mil) have been published. The last two volumes 
(IV- V) are already prepared and will be published 
in June of 1958. 



Res. Bull. Coll. Exp. For. Coll. Agr. 

Hokkaido Univ. 18-1. 1 54. (1956) 



Hiroshima, famous following the first dropping 
of the atomic bomb, was the center of the 
research. It is facing the Inland Sea (Setonaikai) 
and is situated 3424'N., 13227'E. We studied 
a belt which extended from the seaside at Hiro- 
shima to the inland mountains that form the 
backbone of the Chugoku Region. The distance 
is about 42 km, and the rise in vertical elevation 
from sea level to the mountains is 1,346 m. Along 
the transects under consideration, five experi- 
mental localities were studied as follows: 

Name of forest 

Castanopsis cuspi- 

Cinamomum Cam- 

Castanopsis caspi- 

A hies firma-Tsuga 
Fagus crenata 
Quercus crispula 
Tsuga Sieboldii 

Name of 





tion (m) 
















4. a 








, , 





















Abies fir ma 



ibid. 18-2. 53 148. (1957) 

Geobotanically, the Island of Yakushima is 
one of the most important localities in Japan. 
It is extremely interesting, with many endemic 
and rare elements. Yakushima is roughly orbi- 
cular in shape, about 27.1 km long and 26.7 km 
wide, with an area of approximately 544 sq km, 
lying between 13023'-40' East Longitude and 
30 13'- 28' North Latitute. In the central part 
of the island, there are high mountains, such as 
Mt. Miyanoura (1,935 m), Mt. Nagata (1,890m) 
and Mt. Kuromi (1,836 m). All rivers extend 
toward the sea from these central highlands, 
and enroute they often form deep gorges. 

In 1916, the American botanist Dr. H. Wilson 
visited Yakushima and spoke admiringly of the 
beautiful primeval forests ofCryptomeriajaponica. 


Both before and after him, many Japanese bota- 
nists have turned their attention to floristic and 
vegetational studies of the region, among them, 
Profs. G. Koizumi, G. Masamune, S. Hatsushima, 
Drs. M. Kawada, K. Imanishi, Mr. Z. Tashiro 
and so forth. The results of their important 
researches were cited briefly in the paper. The 
studies of Prof. G. Masamune are especially 
important, for he described the flora in excellent 
and precise terms. 

The two islands Yakushima and Tanegashima 
form a special phytogeographical district of the 
East Asiatic Warm Temperate Zone. Tokara 
Strait, lying just to the south of Yakushima 
island, acts as a line of demarcation between this 
zone and the elements at the northern limit of their 
distribution, which perhaps is only natural, con- 
sidering the unique geographical position of the 

But it is quite remarkable that certain Japanese 
elements, which make up a prominent part of the 
flora also have their northern limit of distribution 
in this region, namely: Cephalotaxus drupacea, 
Torreya nucifera, Abies fir ma, Chamaecyparis 
obtusa, Castanca crenata, etc. 

The causes of this peculiar and somewhat 
confined distribution have been attributed to the 
presence of high mountains on Yakushima and 
deep sea separating the islands from the southern 
areas. These geographical barriers may be con- 
sidered to be the primary ecological factors 
operating against a more widespread occurrence 
of the Japanese elements named above. Along 
the transects under consideration, nine localities 
were studied as follows: 

Name of 


Transect Eleva- 
number tion (m) 



Name of forest 

Kandelia Candel 


Nabeyama 3. a 


Kosugidani 4.a 













Ficus Wightiana- 
Ficus retusa 
Machlus Thunder- 


Quercus Wrightii 

Distylium racem- 

Cryptomeria ja- 

6.b 1,100 



Name of Transect Eleva- Name of forest 
locality number tion (m) 

Name of Transect Eleva- Name of forest 


Kosugidani 7.a 960 
7.b 960 

7.c 960 

Mt. Ishizuka 8.a 1,200 
8.b 1,440 


Miyanoura 9. a 1,760 

Abies firma 
Tsuga Sieboldii- 
Cryptomeria ja- 

Tsuga Siebohlii 
Tsuga Sieboldii 
Cryptomeria ja- 
ponica - Trochoden- 
dron aralioides 

Cryptomeria ja- 



ibid. 18-2. 149 208. (1957) 

In some limited localities in Southern Kytishu, 
the forests are characterized by subtropical 
vegetation. Pure forest of Kandelia Candel, Cycas 
revoluta, and Livistona subglohosa, respectively, 
are representative. The latter two forests, occur- 
ring in rather small areas, are found at the points 
of the peninsula protruding into the Pacific Ocean 
washed by warm currents or on the islets sur- 
rounded by warm current. The forest vegetation 
in the low lands of the district under consideration 
predominantly consists of evergreen broad-leaved 
trees. The present study was carried out from the 
seaside to the mountain to heights of about 
1,000 m., ranging from the subtropical forests 
to the temperate needle-leaved forest through the 
evergreen broad-leaved forests. Along the tran- 
sects laid out for study, eight experimental local- 
ities were situated as follows: 

Name of 



Name of forest 



tion (m) 



Kandelia Candel 




Distylium racem- 





11 *1 




Livistona subglob- 





11 11 




Cycas revoluta 








11 11 




Cinnamomum Cam- 





Kirishima 8. a 


tion (m) 

5 Livistona subglob- 

990 Abies firma-Tsuga 


1 ,050 Tsuga Sieboldii 
960 Abies firma 



ibid. 18-2. 149208. (1957) 

The dominant feature of the forest vegetation 
of Southern Shikoku is represented by "pluvii- 
silvae." In several places of the district under 
consideration, the natural physiognomy of the 
forests has fortunately been kept and poses an 
important problem for geobotany in spite of the 
comparatively small areas in question. 

The general arrangement of the forests will be 
explained first. Along the shore, the Pinus Thun- 
bergii forest is commonly developed on sandy 
beaches. On flat terraces, the Pittosporwn Tobira 
forest mixed with Eurya emarginata forms the 
front zone against the sea. It attains a height 
ranging 1-3-5 m influenced by the wind. The 
Camellia ja ponica forest succeeds it and is then 
followed by the Machilus Thunhergii forest 

In the Machilus forest belt, Distylum racetnosum, 
Podocarpus Nagi\ and Elaeocarpus sylvestris are 
sometimes found, forming groves in some areas. 
The pure Quercus phillyraeoides forests are gen- 
erally found on the rocky slope along the shore. 
Nephrolepis cordi folia and Pyrosia Lingua are com- 
monly found in the undcrlayer. The Castanopsis 
cuspidata forest is also found inland, but it has 
lost its primeval physiognomy. Only one place in 
Ikku, near Kochi, a rather young natural forest, 
was selected for ecological analysis. 

The elements of the subtropical forests are well 
represented by Fieus Wightiana and Livistona 
subglobosa. Their areas are very limited. The 
former forest is found in Cape Muroto and Cape 
Ashizuri. The latter is found only in Ashizuri, 
but the question whether it is an invader from 
the south or a remnant is still unsettled. We 
consider it as a vestige. 

Along the transects under consideration, six 
experimental localities were situated as follows: 



Name of Transect 
locality number 


Eleva- Name of forest 
tion (m) 

Ficus Wightiana 
Quercus phillyrae- 

Distylium racemo- 

Castanopsis cuspi- 


Livistona subglob- 

Pittosporum Tob- 



Camellia japonica 

Machilus Thunber- 

Quercus phillyrae- 

Livistona subglob- 




By M. TATEWAKI, etc. 

The Japanese beech, Fagus crenata, is the 
representive tree of the cold temperate forests in 
Japan. Up to the present, the precise study of 
its distribution within the northern limit was 
carried out by Tatewaki in 1948. The geobotani- 
cal study of the Japanese beech forest of the 
district under consideration is a most important 
and interesting problem not only from the view- 
point of geobotany, but also from that of forestry. 
During these ten years, especially during the last 
five years, Tatewaki accompanied by the members 
of his institute, T. Misumi, T. Igarashi, 
S. Watanabe, and S. Kawano, have devoted 
themselves to the study of this question under 

the auspices of the Hakodate Regional Forest 
Office. Along the transects under consideration, 
seven localities were studied, as follows: 










Tosa Ikku 






























,, ,, 












99 19 




sample plot 


Name of Transect 


Name of forest 

locality number 

tion (m) 




Fagus crenata 




99 99 





M ) 

99 99 



91 99 




99 99 





91 19 

Mt, Kariba 



Betula Ermani 

99 99 



Fagus crenata 

99 99 



99 99 

99 99 



99 99 

99 99 








Betula Ermani 

99 99 



99 99 

19 99 



Fagus crenata 

-Betula ermani 

99 99 



Betula Ermani 

-Fagus crenata 





Fagus crenata 




91 99 



11 99 



Mt. Ohira 

4 a 


11 99 



11 99 






11 91 



11 99 



,, ,, 



11 91 







4 j 

















99 99 



99 99 



99 99 



19 99 





99 99 



91 99 



99 99 






University of Florida y Gainesville, Florida ; U.S.A. 


Over the land mass of continental eastern Asia, 
forest communities once flourished in an almost 
unbroken expanse of over 30 degrees of latitude, 
from the tropics to eastern Siberia. Throughout 
the immense geographic range there is a gradual 
displacement of forest components, with resultant 
intergradations and segregations of forest types. 

The forest vegetation of continental eastern 
Asia comprises the following main types 1 : 

(1) The montane coniferous forest formation: 
spruce-fir forest, larch forest; 

(2) The broad-leaved deciduous forest forma- 
tion : the mixed mesophytic forest, the mixed 
northern hardwood forest, the deciduous 
oak forest, birch forest; 

(3) The broad-leaved evergreen forest forma- 
tion: the evergreen oak forest, the rain 
forest, the littoral forest. 

The main types of forest vegetation are arranged 
in approximately the latitudinal and altitudinal 
sequence as presented in the vegetation map. 

Throughout the long geologic past, since the 
advent of angiospermous plants in the late 
Mesozoic Era, there were always areal and com- 
positional differentiations in the vegetation. The 
author assumes that all the natural phenomena in 
geologic time were governed by laws that govern 
the Universe today. The plants of the existing 
vegetation, as were plants throughout the ages, 
are the direct descendants of pre-existing ones 
with which they are genetically most closely 
related. Upon this premise, the developmental 
concept of plant communities used in this paper 
is based. The phylogenetic relationships of the 
components of the plant communities offer the 
most reliable clue in interpretations of develop- 
ment, differentiation and relationship among the 
different forest communities, their present pattern 
of geographic distribution, and their vicissitudes 
during geologic time. 

Evidence afforded by the existing vegetation 
and that of the geologic past suggests that the 
existing forest communities originated from a 
type that resembled the modern rain forest. The 

differentiated, usually simpler and specialized 
types of forest communities, different as they are, 
arc linked with the ancestral type represented in 
the modern rain forest by a continuum of varia- 
tion among phyletic stocks. The modern rain- 
forest has been changed through contraction and 
spcciation among its components, as have other 
forest communities. It still retains the multitude 
of phyletic stocks of which the other forest com- 
munities are composed and to which they are 
linked by a close phylogenetic bond. 

The ancestral community, as exemplified by 
the existing rain forest, is generally represented 
by primitive forms and usually by far more sec- 
tions, genera, and even sub-families. In other 
words, the rain forest is composed of descendants 
from a higher level of plant evolution, and hence 
an older one in time sequence, while the other 
forest communities are composed of later-differ- 
entiated and often specialized forms of the same 
phyletic stocks. 

The compositional and arcal differentiations of 
the forest communities are primarily the results 
of two basic processes: historic contraction and 
regional speciation. They are the manifestation 
of the innate potentialities of all the phyletic 
stocks of forest trees ever evolved on the earth's 
surface. The extreme paucity of components in 
certain simple forests of wide extent, e.g., birch 
forest, larch forest, and the multitude of mono- 
typic groups and ogliotopic genera in the poly- 
phyletic communities, e.g., mixed mesophytic 
forest, furnish evidence of the extent of contrac- 
tion. On the other hand, the clusters of numerous 
closely related species, in both the polyphyletic 
and the monophyletic communities, and the geo- 
graphic varieties, races, and forms recognized 
within population of extensive range bear witness 
to the active process of regional speciation. 

Through contraction, which widened the gaps 
of discontinuity both in character and in range, 
distinct species (such as Populus tremuloides) 
and subgenera (such as the black oak group, 
Erythrobalanus, of North America) and genera 
(such as the evergreen beech group, Nothofagus, 
of the Southern Hemisphere) likewise have 

t Presented by F.E. Egler. 

1 The Forest Types of Continental Eastern Asia I, II. 8th Pacific Science Congress, Manila, Philippines. 



resulted from the species clusters. The differen- 
tiation of subfamilies and families is only the 
prolongation of the process, which in turn trans- 
forms simple types of forest communities into 
polyphyletic types in time to come. 

In the present treatise the segregation and 
development of two main forest types, the 
mixed mesophytic forest and the mixed northern 
hardwood forest, are traced. The closest living 
representatives of these two forest types are found 
in eastern North America. Frequent references 
are made to the forests of eastern North America. 
The author believes that the segregation and devel- 
opment of forest communities, as visualized 
above, is an universal phenomenon and has been 
contemporaneous in time sequence throughout 
the world. But it is more fully expressed in the 
existing vegetation of eastern Asia which extends 
from the tropics to the Arctic and which is the 
greatest repository of Tertiary relics. The main 
types of forest vegetation are not only spatially 
and compositionally differentiated entities, but 
each of them reflects a common level of plant 
evolution. They are linked by a strong bond of 
descent upon which the interpretation proposed 
in this paper is based. 



The mixed mesophytic forest of eastern Asia is 
lloristically the richest in composition of all the 
deciduous broad-leaved forests, and is surpassed 
in complexity only by the rain forest. Not only 
does it include a multitude of tree species, but the 
trees represent a large number of remotely related 
phyletic stocks. It includes also numerous relics 
of an "arcto-Tertiary flora" whose most closely 
related modern species are found in the mixed 
mesophytic forest of North America. The tree 
components of the mixed mesophytic forest of 
eastern Asia include more than 10 genera of 
conifers and over 50 genera of hardwoods ^ever- 
green broad-leaved trees): 


Cunn ingham ia 


Tor re y a 

Broad-leaved trees 

A canlhopanax A cer 

Aphananthe Fagus 


Betula Fraxinus 


Camptotheca Gymnocladus 


Carya Halesia 


*Castanea Hovenia 


* Ccistanopsis Idesia 


Celt is *IHicium 


Cercidiphyllum Juglans 


Cladrastis Kalopanax 


Daphniphy 'Hum Koelreuteria 

* Quercus 

Davidia Liquidatnbar 


Diospyros Liriodendron 


Ehretia *Lithocarpus 


Elaeocarpm Maackia 


Emmenopteris Magnolia 


Eucommia *MangIietia 


Euptelea Meliosma 


Evodia Moms 

The "mixed mesophytic forest climax" of 
eastern North America, as defined by Braun (5, 
40-41), includes 25 species and varieties of dom- 
inant trees in the aboreal layer. They represent 
the following genera: 

















The following genera, which sometimes appear 
in the climax stands, can be added to the above 





All these 30 genera are represented in the mixed 
mesophytic forest of the Yangtze Valley, except 
for Oxydendron, Plalanus, and Robinia. 

Braun (4* 5) maintains that the mixed meso- 
phytic association of eastern North America is the 
"lineal descendant" and persisting remnant of the 
"undifferentiated forest of the Tertiary," and the 
most complex and oldest association of the 
deciduous forest formation. It occupies a central 
position in the deciduous forest as a whole; and 
from it or its ancestral progenitor (the mixed 
Tertiary forest) all other climaxes of the decidu- 
ous forest have arisen. 

How satisfactory this center of radiation theory 
for the mixed mesophytic forest is, will be verified 



by further studies of my American colleagues. 
It is known, however, that the Tertiary forest of 
the middle latitudes of eastern North America was 
far richer in tree components than the existing 
mixed mesophytic forest. It included the follow- 
ing genera that do not exist in the present forest 
(Wilcox flora, Eocene) : 

* Acacia 






Ar Wear pus 


Psycho tria 


* Dalbergia 

* Sap Indus 





* Engelhardtia 



* Eugenia 








* Cassis 







The fossils assigned to " Dryophyllwn" (3) are, 
according to Sharp (18), more closely related to 
modern Mexican oaks. Representatives of these 
genera now exist in the broad-leaved evergreen 
forest region of eastern Asia, and the majority of 
them (*) still can be found in the southern part of 
eastern North America south of the mixed meso- 
phytic forest region, mostly in Florida and 
Mexico (18). 

The present mixed mesophytic forest of eastern 
North America is primarily a deciduous commu- 
nity. If it is the lineal descendant of the Tertiary 
forest, as suggested by Braun, the evidence indi- 
cates that the existing forest has been changed 
considerably from its progenitor. 


The most obvious change in the composition 
of the mixed mesophytic forest has been the 
nearly total (as in eastern North America) or 
partial (as in eastern Asia) elimination of ever- 
green broad-leaved trees. This change resulted 
in a transformation into an essentially deciduous 
community, such as the mixed mesophytic forest 
of today, from a forest community that was 
dominated by, or at least, included a considerable 
portion of evergreen broad-leaved trees in the 
arboreal layer. The modern equivalents of these 
Tertiary evergreen broad-leaved trees are mostly 
associated with the rain forest of today. 

This trend, as indicated by fossil evidence, is 
also suggested by the components of the existing 
forest. The fact that the greatest concentration of 
epibiotics is in the mixed mesophytic forest is of 

special significance as a clue to the historic 
contraction of this community. 

Most of the epibiotics of the mixed mesophy- 
tic community are represented by monotypic gen- 
era, or by monotypic family, such as: 







Dipt crania 

















The above-mentioned plants include only those 
epibiotics of regional distribution which charac- 
terize the geographic intergradations within the 
mixed mesophytic community. The epibiotics 
arc not all necessarily restricted in range, however. 
The following include those that are distributed 
in the entire region of the mixed mesophytic forest 
and beyond. They are either monotypic groups 
or oligotypic "polytopic genera," i.e., small genera 
composed of only a few geographically disjunc- 
tive species, usually isolated in the remote parts of 
the world. 

(1) Monotypic families: In addition to Amento- 
taxaceae, Bretschneideraccae, Cercidiphyllaceae, 
Metasequoiaceae, and Rhoipteleaceae mentioned 
above, are: Ginkgoaceae (Ginkgo) , endemic to the 
mixed mesophytic region of the Yangtze Valley, 
and Eucommiaceae ( Eucommia), Yangtze Valley 
and the transitional zone of the Northern Pro- 

(2) Monotypic genera: In addition to those 
mentioned in the last section (Davidia, Nothot- 
axus, Pseudolarix, Taiwania, Tetracentron), are 
the following: 




Eusc aphis 








(3) Poly topic genera: The following include those 
polytopic genera that now occur in more or less 
the entire range of the mixed mesophytic commu- 
nity. Those genera that have persisted only in 
certain parts of this region were discussed in the 
preceding section. 



Aphananthe Eastern Asia and Australia 3-4 
species, China 1 

Buckleya Continental eastern Asia 3, east- 
ern North America 1 

Chionanthus Continental eastern Asia 1, east- 
ern North America 1 

Cladrastis Continental eastern Asia 2, Japan 
1 , eastern North America 1 

Cunninghamia Yangtze Valley and farther 
south 1, Taiwan 1 

Gymnocladus Continental eastern Asia 1, east- 
ern North America 1 

Liquidamhar Eastern Asia 1, western Asia 1, 
North and Central America 1 

Liriodendron Continental eastern Asia 1, east- 
ern North America 1 

Nyssa Continental eastern Asia 1, east- 

ern North America 4, Himalaya 
to Java 1 

Pseudotsuga Continental eastern Asia 2, west- 
ern North America 3-4 

Sassafras Continental eastern Asia 1 , Tai- 
wan 1, eastern North America 1 

The multitude of relics in the mixed mesophy- 
tic forest is in sharp contrast to their extreme 
paucity in the areas just to the north, which 
are occupied by plant communities developed 
over a new land surface and in a new habitat 
resulting from climatic changes in relatively recent 
geologic history. This fact seems to indicate 
that the mixed mesophytic forest is situated at the 
periphery of the selective pressure which has 
been effective enough to decimate part of the 
population, and yet favorable enough for the 
survival of the rest- a fact which possibly explains 
the preservation of the numerous ancient relics 
and the extremely rich iloristic composition of 
the mixed mesophytic forest. 

Braun (5) aptly observed that the comparable 
community of North America is "the most 
complex and the oldest association of the Deci- 
duous Forest Formation." The complexity of 
this type of forest, however, is not due simply 
to the multiplicity of tree species. Strikingly 
indeed, it is composed of many t4 one-of-a-kind" 
elements. The 60 species of crown trees that are 
found throughout the entire range of the mixed 
mesophytic community represent no less than 
50 genera. This fact and the presence of the 
numerous monotypic groups and polytopic genera 
suggest that the mixed mesophytic type, complex 
as it seems to be, is but an impoverished relic of a 
floristically much richer community. 



The original composition before the impover- 
ishment included more genera that are related 
to the trees of the rain forest, as suggested by the 
history of the mixed mesophytic forest of eastern 
North America. In the present forest of eastern 
Asia the rare occurrence of such genera as 
Podocarpus, Torrcya, lllicium, Phoebe, Manglietia, 
and evergreen oaks (Quercus, Castanopsis, Pas- 
ania) suggests similar conditions. 

The comparative morphology of angiosperms 
(19, 1 ) suggests that the deciduous trees in general 
arc derived from evergreen stocks which are close 
to the more primitive forms. The major compo- 
nents of the mixed mesophytic forest, which are 
naturally all deciduous trees, are nearly all repre- 
sented in the rain forest by evergreen trees of 
closely related genera or species. In fact, most of 
the trees of the mixed mesophytic forest as listed 
below are deciduous representatives of phyletic 
stocks which are predominantly evergreen, viz: 

( Diospyros) 

( Elaeocarpus ) 

( Daphniph vllum, Mal/otus) 

(Poliothyrsis, Idesia) 

(Llquidambar , Forlunearia, 


(Sassafras, Under a, Lit sea) 
( 1. iriodendron , Telracen iron ) 
( Morus) 
( Da vi did) 

(Chionanthus, Fraxinus) 
( Emmenopterys) 
(Evodia, Phvllodendron ) 
(Buckleya) ' 
( Koelreuteria) 
(Tapis c\a, Euscaplus) 
(Halesia, Sinojackia, Huo- 

dendron, Rehderodendron, 


Two species of the genus Castanea in the mixed 
mesophytic forest are evergreen. Even Acer, 
which is represented by over 50 species of decidu- 
ous trees in eastern Asia, and the Boraginaceae, 
which is essentially a family of herbaceous plants 
of the arid regions, are both represented by decidu- 
ous trees (deciduous Acer and Ehretia) in the 
mixed mesophytic forest, but by evergreen trees 
in the tropics (section of entire leaved maples, 
Cordia Ehretia, and Tournefortia). The woody 
boraginaceous plants represent the primitive 
section of the family (9) . 

The closest kin of the deciduous trees of the 
mixed mesophytic forest can nearly all be traced 



















to the rain forests of the tropics, which usually 
include more tribes, genera, and sections of the 
phyletic stocks to which the deciduous trees of 
the mixed mesophytic forest are related in a 
continuum of variation. The weight of evidence 
indicates that the mixed mesophytic forest has 
emerged from the northern fringe of a type of 
evergreen broad-leaved forest similar to the 
present rain forest. The evergreen components 
were gradually eliminated, probably in late 
Tertiary. Through further contraction and 
speciation the multitude of relics and numerous 
closely allied species of certain large genera of 
deciduous trees which characterize the present 
mixed mesophytic forest were differentiated. 


The mixed northern hardwoods forest, in conti- 
nental eastern Asia is situated between the boreal 
coniferous forest farther to the north and the 
deciduous oak forest of the relatively arid North- 
ern Provinces, which in turn separates the mixed 
northern hardwoods from the mixed mesophy- 
tic forest of the Yangtze Valley. Despite the 
discontinuity, the mixed northern hardwood 
forest is closely related to the mixed mesophytic 
forest in composition. It appears to be a severely 
depauperized form of the mixed mesophytic 


The primary components of the mixed northern 
hardwoods are Acer (8 species), Tilia (5 species), 
and Betuht (9 species), intermixed with consider- 
able proportions of white pine (Pinus koraiensis), 
Quercus, Fraxinus, Juglans* Maackia, Phelloden- 
dron, and Ulnms. This forest also includes, as 
minor components, trees of the following genera: 
Alnus Mains Pyrus 

Carpinus Moms Salix 

Celtis Populus Sorbus 

Kalopanax Prunus Zelkova 

Taxus, Magnolia, and shrubby Cornus and Lindera 
also occur as rare accessories. 

In comparison with all the other types of deci- 
duous broad-leaved forests, the mixed northern 
hardwoods is, next to the mixed mesophytic, the 
richest in composition. All the above mentioned 
26 genera are common to both types. However, 
the specific representative of these genera usually 
are not the same. Furthermore, there are many 
more genera (over 40) of evergreen and deciduous 
trees among the primary and minor components 
of the mixed mesophytic forest that are not repre- 

sented now in any other type of deciduous forest. 
But some of them are known to have occurred 
in the northern regions in the Upper Tertiary. 
This fact shows the gradual retraction of the range 
of certain "southern" genera and the depaupe- 
rization in components of the northern lobe of a 
more extensive mixed mesophytic forest which 
once flourished in the northern region. This 
process resulted in the simpler type of forest 
community of deciduous broad-leaved trees now 
designated as the mixed northern hardwood forest. 
All the genera of the forest components, without 
exception, are included in the present mixed 
mesophytic forest, and some of the eliminated 
components are preserved in the Tertiary fossils 
of this region. 

With the exception of Tsuga and Fagus, all the 
genera of the primary forest components of this 
community in North America are represented in 
the mixed northern hardwood forest of eastern 
Asia, which is somewhat richer in composition 
than its North American counterpart. Common 
forest components such as Phellodendron, Maac- 
kia, and Zelkova and a common climber, Actinidia, 
are not known in North America. Schizandra 
is represented in North America only in the South. 
Recent interspecific hybridization (8) between the 
white pine of the Northeastern Provinces of 
China (Pinus koraiensis) and the eastern white 
pine of North America (Pinus strobus), both of 
which are important components of the respec- 
tive mixed northern hardwoods, gives almost 
100% fertility. The striking compatibility of the 
two vicarious species furnishes genetic evidence 
for the close affinity of the two similar but widely 
disjunct forest communities. 


An entirely different interpretation of the 
formation and the present distribution has been 
proposed for the mixed northern hardwood forest 
of eastern North America. According to Braun 
(5, p. 533), the vegetation of the mixed northern 
hardwoods is "the result of post- Wisconsin migra- 
tions, which brought about an expansion from the 
unglaciated Allegheny Plateau and northern 
Allegheny mountains (physiographic sections of 
Appalachian Plateaus) without pronounced modi- 
fication of type. The climax elements are almost 
entirely a result of these expanding migrations." 

In the interpretation of the present distribution 
of the deciduous forests of eastern North America, 
great pains were taken to correlate the physio- 
graphic history with forest development, particu- 
larly the Pleistocene glacial boundary. The ice 



advanced across the Appalachian Plateau, the late 
Tertiary haven of the mixed forest. From fossil 
evidence, Braun (5, pp. 512-513) deduced that 
"nowhere far beyond the glacial boundary was 
climate during the glacial stages sufficiently severe 
to displace occupying vegetation," and this 
ensured continuous occupancy by the late Tertiary 
mixed forest, the progenitor of the present mixed 
mesophytic forest, on the unglaciated part of the 
Appalachian Plateau and a small isolated ungla- 
ciated area (the Reading Prong and the New 
Jersey Highlands) farther east. Braun (5, p. 529) 
maintains "Continuity of occupation since early 
Tertiary time accounts for the antiquity of its 
vegetation ; lack of extreme changes, for the con- 
ti nuance of one climax type; and a consistently 
humid climate (with rare lapse to subhumid) 
for the prevalence of this climax/' The explana- 
tion of the present forest distribution as the result 
of post-Pleistocene expansion and segregation of 
the vegetation from the unglaciated areas to the 
glaciated areas where the former vegetation was 
removed by the advancing ice, is plausible enough. 


However, it does not necessitate the continuous 
occupancy of the late Tertiary mixed forest not 
far from the front of the continental ice sheet, an 
assumption that is still open to debate. Pcrigla- 
cial phenomena such as block-fields and thick 
surficial deposits moved by soli-fluction probably 
of Wisconsin age are found in the higher parts of 
the Appalachian Highlands as far south as the 
Great Smoky Mountains (6,600 ft, circ. Lat. 
35 N.) where the cove hardwoods type (circ. 
3,000 - 3,500 ft) is considered as the most typical 
of the mixed mesophytic communities. At the 
present time, areas in the subarctic or in high 
mountains where similar periglacial phenomena 
are actively forming are all essentially treeless. 
By analogy, Denny reasons that the Appalachian 
Highlands were likewise essentially devoid of 
forest when such periglacial phenomena were in 
action ( 1 ). 

This inference does not entirely preclude the 
continuous occupancy of the late Tertiary mixed 
forest in the unglaciated areas still farther away 
from the glacial boundaries. Tt does indicate, 
however, that the Tertiary mixed forest of eastern 
North America, like the similar type of eastern 
Asia, has undergone severe displacement, espe- 
cially in regions close to glaciations. It also 
suggests that to a considerable extent the present 
mixed mesophytic forest is the result of post- 

Pleistocene migrations from areas farther to the 

There are indisputable evidences of Quater- 
nary glaciations in the mixed mesophytic forest 
region of eastern Asia (13, 14, 15, 2, 21) . Glaci- 
ated areas include: Lushan (1,480 m, Lat. 
2930' N.), Chiuhuashan (900 m, Lat. 3032' N.), 
Tienmushan (Tienmongshan, 1,547 m, Lat. 
3025' N.), and Huangshan (1,700 m, Lat. 30 
10'N.). These mountains are situated in the 
southern part of the Lower Yangtze Valley, not 
too far from the coast. 

The Lower Yangtze Valley, which is now occu- 
pied by mixed mesophytic forest, was glaciated 
not once but three times. In the Poyang glacia- 
tion (the oldest and also the largest) and the next, 
the Taku glaciation, piedmont glaciers were 
formed. The third, the Lushan glaciation, pro- 
duced only small glaciers in the higher mountains. 
These glaciations were separated by interglacial 
periods of genial or even subtropical climate, 
resulting in the high oxidation of the ferruginous 
soil and incipient lateritization of the boulder-clay- 
like deposit and the Red Loam (76, chapter on 
Pleistocene climate). 

Despite the successive glaciations, the existing 
vegetation of the above-mentioned glaciated 
mountains, as evidenced by the preserved natural 
forests, is of the mixed mesophytic type which 
includes not only nearly all the tree genera of the 
mixed mesophytic forest of the unglaciated Appa- 
lachian Plateau, but also survivals from those 
genera that are known to have occurred in the 
late Tertiary mixed forest of eastern North 
America and to have perished subsequently. 

There are distinctions between forest types and 
forest regions. The mixed forest of the Miocene 
whose components were almost identical with 
those of the present mixed mesophytic forest but 
which occurred more than 500 kilometers north 
of the northern limit of the present type, has been 
replaced by the present deciduous oak forest. 
It is then reasonable to infer that in the Quater- 
nary glaciations the areas that are now occupied 
by mixed mesophytic forest, but were at that time 
glaciated or at least exposed to periglacial condi- 
tions, could not possibly have been occupied by a 
type that was similar to the present mixed meso- 
phytic forest. The presence of the mixed meso- 
phytic forest, no matter how complex or ancient 
the flora may be, does not imply that the area has 
never been glaciated nor does it necessitate the 
continuous occupancy of this type in that parti- 
cular area. 




Nor is it necessary to assume the total elimina- 
tion of forest communities and to preclude the 
possible persistence of a depauperized form of the 
late Tertiary mixed forest farther to the north, 
particularly in the maritime regions where the 
influence of warm currents prevailed. The mixed 
northern hardwoods forest, Braun maintains, was 
almost entirely a result of post-Wisconsin migra- 
tion so far as the climax elements are concerned. 
To her disadvantage, the forest flora of the 
deciduous forests of eastern North America with 
which she was dealing, rich as it is, is relatively 
simple in comparison with that of eastern Asia. 
The geographic intergradations in the whole range 
of the American deciduous forests are not well 
expressed by the regional speciation of their 
components. Of the 25 species and varieties listed 
as "dominant trees of the arboreal layer" of the 
mixed mesophytic forest of eastern North Ame- 
rica, all are represented in other "climaxes of 
deciduous forest." Gymnocladus, Halesia, Liqui- 
dambar, and Oxydendron are perhaps the only 
tree genera of the mixed mesophytic forest that are 
not represented in the deciduous forests of the 
North. Not only are most of the genera in com- 
mon, but their representatives in the primary 
components of both the northern hardwoods 
and the mixed mesophytic forest are of the same 
species, which extends in a continuous range. This 
fact seems to have led Braun to the conclusion 
that all the other deciduous forest climaxes have 
arisen from the centrally located "undifferentiated 
mixed mesophytic forest," supposedly the oldest 
association of the deciduous forest formation 
preserved in the Appalachian Plateau. 

In a forest flora like this, evidence of geo- 
graphic differentiations could be found at the 
subspecific level, e.g., the local races and eco- 
types. Recent studies of the American beech 
(6), ash (22, 23) walnut (24), maple (10), hem- 
lock (17) demonstrate that forest trees are made 
up of a number of more or less distinct types with 
well-differentiated ranges. 

In the deciduous forest communities of eastern 
Asia, on the other hand, geographic differentia- 
tions are well expressed on the species level. The 
mixed mesophytic forest and the mixed northern 
hardwoods in particular, are not only richer in 
components, but the tree genera of the mixed 
northern hardwoods that are in common with the 
mixed mesophytic forest are represented mostly 
by different species. In other words, distinct re- 
gional speciation has taken place along with the 

differentiation of forest types. The interpreta- 
tion advanced by Braun of the mixed northern 
hardwoods of eastern North America cannot 
satisfactorily explain this phenomenon. The 
mixed northern hardwoods forest of eastern Asia 
cannot possibly be entirely the result of post- 
Pleistocene expanding migrations from un- 
glaciated areas south of the ice front : 

(1) There are only a few species of trees in the 
mixed northern hardwoods that are connected 
with the mixed mesophytic forest in a more or less 
continuous range. Most of the other non-endemic 
species of the primary components are generally 
restricted to the north and rarely occur south of 
the deciduous oak forest region. These few wide- 
ranging trees occur both in the mixed mesophytic 
forest of the south (MM), the deciduous oak 
forest of the northern Provinces (N), and the 
mixed northern hardwoods of the Northeastern 
Provinces (NE). Some of them extend as far as 
Japan (J), Korea (K), eastern Siberia (S), and 
Sakhalin (SK). This group of wide-ranging trees 
includes the following: 

Acer mono (A. pic turn var. parviflorum) (MM, 
N, NE, K, J) 

Betula japonica (B. mandshuricd) (MM, N, K, 
J, S, SK) 

Carpinm cordata (MM, N, NE, K, J, S) 

Fr ax inns chinensis (MM, N, NE) 

Populus iremula var. davidiana (MM, N, NE) 

Ulmus japonica (MM, N, NE, K,J) 

V.pumila (MM, N, NE, S) 

A number of lesser plants can be added to this 
list, e.g., Schizandra chinensis (MM, N, NE, K, J, 
S) and Actinidia arguta (MM, N, NE, K, J, S). 
The true mistletoe, Viscum album, unlike the 
American form, Phoradendron Flavescans, extends 
from the mesophytic forest to the boreal regions 
(Siberia) with a total disregard for glacial bound- 

(2) The majority of the primary components 
of the mixed northern hardwoods are tree species 
of a northern range which rarely if ever appear in 
the mixed mesophytic forest region. A number of 
these northern trees are endemic to the Northeast- 
ern Provinces and the adjacent part of Korea 
where this type of forest is extensive, or restricted 
to the general region of the Northeastern Pro- 
vinces (NE), Korea (K), Japan (J), and eastern 
Siberia (S). Some of them extend even to 
Sakhalin (SK). This group of trees includes the 

Acer barbinerve (NE, K) 
A. Mandshurica (NE, K) 
A. pseudo-sieboldianum (NE, K) 



A. te%mentosum (NE, K) 

A. triflorum (NE, K) 
Alnus hirsuta (NE, K, J, S) 
Be tula cos tat a (NE, K, S) 

B. er/wfl/i/ii (NE, K, J, S) 
B. platvphvlla (NE, K, S) 
B. Schmidtii (NE, K, J, S) 
Fraxinus mandshurica (NE, K, J) 
Maackia amurensis (NE, K, J) 
Phellodendron amurensis (NE, K, S, SK) 
Sorbus amurensis (NE, K, S) 

7Y//0 amurensis (NE, K, S) 

7. megaphylla (NE) 

Pinus koraiensi\ (NE, K, J) 

Ttfjotf cuspidata (NE, K, J, S, SK) 

Magnolia sieboldii (NE, J, K, S), Cornus alba 
(NE, K, S), K/Vw amurensis (NE, K, S), and a 
number of shrubs and lesser plants can be added 
to this list. None of them extends to the mixed 
mesophytic forest region from which the mixed 
northern hardwoods is separated by the decidu- 
ous oak forest region. 

(3) The discontinuity of the two allied forest 
types, the mixed northern hardwoods and the 
mixed mesophytic, is paralleled by the similar 
disjunct occurrence of the following genera: 

Alnus (MM, NE, K, J, S) 
Lindera(MM, NE, K, J) 
Maackia (MM, NE, K, J) 
Phellodendron (MM, NE, K, J) 
Taxus(MM, NE, K, J, S) 

The following plants occur in both the mixed 
northern hardwoods and the mixed mesophytic 
forest, but not in the intervening area: 

Acer caudatum (Himalaya, MM) 

A. Caudatum var. ukwunduense (A. ukurun- 

duense) (NE, K, J) 
A. ginnala (MM, NE, K, J) 
Actinidia kohmicta (MM, NE, J, K, S) 
A. polygama (MM, NE, K, J) 
Lindera mollis (MM, NE) 
Kalopanax pic turn (MM, NE, K, J) 
Phellodendron sachalinense (MM, K, J, SK) 
Samp locos paniculata (MM, NE, J) 

The distribution patterns of these three groups 
indicate progressive contraction in an extensive 
mixed mesophytic type of forest which once 
extended in a continuous range to the present 
mixed northern hardwoods region of the North- 
eastern Provinces and the adjacent maritime areas 
of eastern Siberia. This disjunction of some of 

the components probably is due to the increased 
continentality in the intervening area. Tertiary 
fossil deposits show the actual existence of the 
pre-Quaternary continuity of this type of mixed 
forest. Trees almost identical with living ones 
in the present mixed northern hardwoods and the 
mixed mesophytic forest once occupied the inter- 
vening area. 

In addition to the compositional affinities, the 
pre-Pleistocene continuity of a mixed hardwood 
forest community, and the present disjunct 
range of those components that are common to 
both the mixed northern hardwoods and the mixed 
mesophytic forest, favor the explanation that the 
present mixed northern hardwood forest is a 
depauperized form of a pre-Pleistocene mixed 
hardwood forest community that was similar 
to the present mixed mesophytic forest. The 
prc-Pleistocenc community extended over a 
continuous wide range with an essentially homo- 
geneous composition but with well-developed 
geographic differentiations of tree species. Region- 
al speciation perhaps resulted in the numerous 
endemic species in the subsequently disconnected 
and depauperized forest communities, although 
all the genera of these endemic species and, in 
fact, all the genera, without exception, of the 
primary components of the present mixed north- 
ern hardwoods are represented in the present 
mixed mesophytic forest. 

The presence of the endemic species, parti- 
cularly those that are restricted to this region 
and the adjacent Korea- Japan-eastern Siberia- 
Sakhalin areas, precludes a post-Pleistocene 
origin of the mixed northern hardwood forest. 
If the present mixed northern hardwoods forest 
were the result of post-Pleistocene expanding 
migration from unglaciated areas, as proposed by 
Braun for the mixed northern hardwood forest 
of eastern North America, then the amelioration 
of climate should be such that the entire region, 
including the intervening area, would be suffi- 
ciently favorable for the continuous extension of 
a type of mixed hardwood community that 
connected both the mixed northern hardwoods 
and the mixed mesophytic forest in gradual 

The establishment of a forest community 
including such disjunct plants as Lindera, Symplo- 
cos, Taxus, Maackia, Phellodendron, and a few 
species of maples in the intervening area a 
condition which the theory of post-Pleistocene 
expanding migration requires would necessitate 
a substantial increase in precipitation and a 
milder climate than the prevailing one. 



If it is assumed that the mixed northern hard- 
wood forest of the entire range is of post-Pleisto- 
cene origin, then the implication is that the com- 
ponents common to this region have not only 
migrated to Siberia but have also crossed a land- 
bridge to Japan and Sakhalin. A direct land 
connection between the mainland and Japan and 
Sakhalin Island was possible at the height of the 
Ice Age when the sea level was sufficiently 
lowered. However, such a land connection could 
not have been in existence when the climate 
became sufficiently warm and humid to effect 
the supposedly post-Pleistocene expanding mi- 

On the other hand, if it is assumed that only the 
mixed northern hardwood forest of the mainland, 
i.e., the Northeastern Provinces (Manchuria) 
and eastern Siberia, is the result of post-Pleis- 
tocene expanding migration, and that similar 
forests in Japan and Sakhalin are relics of the 
Tertiary, then the persistence of these forests in 
Japan, and particularly in Sakhalin which is 
situated farther north and is more rigorus in 
climate, serves only to prove the possible survival 
of the mixed northern hardwood forest. This 
forest includes a number of the same species of 
trees in at least the maritime regions of the 
continent just across the sea. 

Pleistocene fossils also indicate that the present 
mixed northern hardwood forest is the depauper- 
ized form of a Tertiary mixed hardwood forest 
rather than that it is of post-Pleistocene origin. 
Zelkova (MM, N, NE, K, J), now extinct in Sibe- 
ria, was growing with Ginkgo in the early Pleis- 
tocene on the Amur River, the frontier between 
the Northeastern Provinces and Siberia (11). 
Juglans mandshurica (N, NE, K, S) now occurs 
on the mainland only, but was found in the 
Pleistocene deposits of Japan (12). The fossil 
record shows that a progressive reduction in 
components and shrinkage in their range had 
taken place. The continuous occurrence of a 
mixed deciduous hardward community which 
was slightly richer in composition and which had 
a more northerly limit than the present mixed 
northern hardwood forest is evidenced by fossil 
remains to have occurred as late as the early part 
of Quaternary. 


So far, the evidence used in the discussion of 
forest development has been limited as much as 
possible to the primary forest constituents, with 
only occasional excursions into the lesser growth, 
so that the main thesis can be presented with 

clarity and effect, without unnecessary reference to 
long lists of plants which have already been a 
burden to the reader. In the mixed northern 
hardwoods forest now under discussion, certain 
lower plants offer complementary evidence for 
the continuous persistence of the type during 
the Pleistocene in this general region. The rare 
occurrence of Hymenophyllum and Trichomanes 
in the present mixed northern hardwoods and 
coniferous forests on the frontier between Siberia 
and the Northeastern Provinces is of special 

Hymenophyllum and Trichomanes, are two 
genera (sen. Lat.) of Hymenophyllaceac which 
include 400-500 species of filmy ferns, mostly in 
shaded habitats of the humid parts of the tropics. 
In North America, one species (Trichomanes 
hoschianum) reaches as far north as Kentucky 
and Illinois. Three species occur in Europe. 
The numerous species of Hymenophyllum and 
Trichomanes of eastern Asia are concentrated in 
the evergreen forest region. They occur only 
occasionally in the mixed mesophytic forest, and 
have been observed in the exceedingly wet valleys 
of western Yunnan in Abies-Picea- deciduous 
hardwood forests. Over the great distance 
between the Yangtze Valley and the Siberian 
frontier, the Hymenophyllaceac have never been 
observed nor are they likely to be found. They 
sometimes reach high latitudes in both hemi- 
spheres, but only under the conditions of maritime 
climate, e.g., Hymenophyllum tunbridgense on 
the Faroe Islands (Lat. 63' N.) in the North 

Kryshtofovich (II, 12) and Vorobiov (20) 
reported the discovery of Trichomanes parvulum 
and Hymenophyllum \\rightii in the Suchan River 
basin of eastern Siberia (Ussuri), close to the 
Northeastern Provinces, a considerable distance 
from the coast. Hymenophyllum wrightii was 
previously known from Sakhalin and Japan (as 
far north as Hokkaido), but not from Korea 
proper except on Quclpart Island. 

In eastern Siberia, Hymenophyllum and Tricho- 
manes grow together on a cliff within a virgin 
forest of Picea ajanesis, Abies nephorlepis, and 
Taxus cuspidata. A rare fern, Pleurosoriopsis 
makinoi, is found nearby. In this general region, 
Vitis amurensis, Schizandra chinensis, Actinidia 
spp., Phellodendron amurense, and Juglans mand- 
shurica may still be found. Kryshtofovich 
concluded that they were survivals of the Tertiary 

The forest around the growth of Hymenophyl- 
lum on Sakhalin consists mainly of Picea ajanensis, 



and Abies sachalinensis, with admixtures of Betula 
japonica, Taxus cuspidata, and Euonymus sachali- 
nensis. The soil is thickly covered with Osmunda 
cinnamomea and Athyrium pterorhachis. In 
places, Ilex rugosa covers the soils with a thick 

The forests of western Yunnan in which 
hymenophyllaceous ferns occur are thousands 
of kilometers away from the Siberian coast; 
nevertheless, the composition is essentially the 
same, including Abies delavayi, Picea complanata, 
Taxus chinensis, Betula caudate, B. albosinensis, 
Sorbus spp., and Magnolia globosa, A variety of 
Betula japonica (var. szechuanica) is known to 
this region. The consistency of the composition 
of the kind of forest in which hymenophyllaceous 
ferns are associated in such widely disjunct areas 
suggests the probable conditions of the relic forest 
that survived the Pleistocene Epoch in the mari- 
time regions of eastern Siberia and adjacent 

In the above discussions on the segregation and 

development of forest communities in continental 
eastern Asia, frequent references have been made 
to the similar type of forests in eastern North 
America. This comparison deserves critical 
examination because two diametrically different 
conclusions were reached from observations of 
two forests that are exceedingly alike in com- 
position, developed under essentially similar 
habitat conditions, and situated in approximately 
the same latitudinal ranges. 

The differences in opinion do not imply the 
author's approval or disapproval of the theory 
advanced by his American colleague. The author 
realizes that the forest communities of the two 
land masses are similar but not the same. How- 
ever, the author does believe that the basic pro- 
cesses that shaped the existing vegetation, as 
revealed by available evidence and presented in 
this treatise, are universal and contemporaneous 
in the general time sequence, but the extent of 
contraction and speciation has involved regional 






(1) Bailey, I.W. and E.W. Sinnott, 1916, The 

Climatic Distribution of Certain Types 
of Angiosperm Leaves, Am. J. Dot., 3: 


(2) Harbour, G.B., 1934, Analysis of the Lushan 

Glaciation Problem, GeoL Soc. China 
Bull., 13: 647-656. 

(3) Berry, E.W., 1930, Revision of Lower 

Eocene Wilcox Flora, U.S.G.S. Prof. 
Paper 156: 1-196. 

(4) Braun, E.L., 1947, Development of the Deci- 

duous Forests of Eastern North Ameri- 
ca, Ecol Monograph, 17: 211-219. 

(5) Braun, E.L., 1950, Deciduous Forests of 

Eastern North America, Blackisten Co., 

(6) Camp, W.H., 1951, A Biogeographic and 

Paragenetic Analysis of the American 
Beech (Fagus), Yearbook, Am. Phislo- 
sophical Soc., Philadelphia 1950: 166- 

(7) Denny, C.S., 1951, Pleistocene Frost Action 

near the Border of the Wisconsin 
Drift in Pennsylvania, Ohio J. ScL, 51 
(3): 116-125. 

(8) Johnson, A. G., 1952, Personal Communica- 


(9) Johnston, T.M., 1951, Studies in the Bora- 

ginaceae XX. Representatives of Three 
Subfamilies in Eastern Asia (Cordioi- 
deae, Ehretioideae, and Heliotropioi- 
deae), /. Am. Arb., 32 (1): 1-26. 

(10) Kriebel, H.B., 1956, Patterns of Genetic 

Variation in Sugar Maple (unpublished 
Doctoral Dissertation). 

(11) Kryshtofovich, A.N., 1 930, Hemenophyllum 

wrightii V.D.B. on the Sakhalin Isl., 
Bull, dn Jard. Bot. Principal de L'UESS 
f., XXIX, L/vr., 3-4: 1412. 

(12) Kryshtofovich, A.N., 1935, Hemenophyllum 

and Trichomanes the Ussuriland of the 
USSR. Sunyatsenia, 3: 22-25. 

(13) Lee, J.S., 1933, Quaternary Glaciation in 

the Yangtze Valley, GeoL Soc. China 
Bull., 8: 15-44. 

(14) Lee, J.S., 1934, Data relating to the study 

of the Problem of Glaciation in the lower 
Yangtze Valley, GeoL Soc. China Bull., 

(15) Lee, J.S., 1936, Confirmatory evidence of 

Pleistocene Glaciation from the Huang- 
shan, Southern Anhui, GeoL Soc. 
China Bull., 15:279-290. 

(16) Lee, J.S., 1939, The Geology of China, Lon- 


(17) Olson, J.S. and H., Hiensteadt, 1957, Pho- 

toperiod and chilling control Growth of 
Hemlock, Sciences, 125 (3246): 492-494. 

(18) Sharp, A.J., 1951, The Relation of the 

Eocene Wilcox Flora to Some Modern 
Floras, Evolution, 5(1): 1-5. 

(19) Sinnott, E.W. and Bailey, I.W., 1915, Inves- 

tigations on the Phylogeny of the Augio- 
sperms 5: Foliar Evidence as to the 
Ancestry and Early Climatic Environ- 
ment of the Augiosperms, Am. J. BoL, 
2: 1-22. 

(20) Vorobiov, P.P., 1933, A Discovery of 

Hemenophyllum wrightii V.D.B. in the 
Suchnan Region of the District, Bull. 
Far East Branch, A cad. Set. ISSR, 1-2-2: 

(21) Wissmann, H. Von., 1937, The Pleistocene 

Glaciation in China, GeoL Soc. China 
Bull., 17: 145-168. 

(22) Wright, J.A., 1944 A, Genotypic Variation 

in White Ash, J. Forestry, 42 (7): 

(23) Wright, J.A., 1944B, Ecotypic Differentia- 

tion in Red Ash, J. Forestry, 42 (8): 

(24) Wright, J.A., 1954, Preliminary Report on a 

Study of Races in Black Walnut, J. 
Forestry, 52 (9): 673-675. 





Pacific Vegetation Project, Care of National Research Council, Washington, D.C., U.S.A. 

Coral atolls are flat islands and reefs of lime- 
stone of organic origin lying only slightly above 
sea level and not in immediate proximity to higher 
land. They are in the form of groups of small 
islands or islets on reefs that enclose shallow 
bodies of sea water called lagoons. Atolls are 
very common in tropical seas, especially the 
central and western Pacific, central and western 
Indian Ocean, and in the Caribbean region. 
They are almost lacking in the eastern Pacific, the 
Atlantic, and the eastern Indian Ocean. 

Because of the relative uniformity of their sub- 
stratum, topography, and temperature, their 
vegetation is rather simple, though by no means 
uniform. Their wide diversity of rainfall and 
geographical position contributes to a degree of 
variation in vegetation that would perhaps not 
otherwise be expected. 

The atoll habitat seems to be a very young one, 
geologically speaking, though it may well have 
existed in something like its present form many 
times in the past. The occurrence after the last 
glaciation of a warm dry period or "climatic op- 
timum" would logically have brought about 
a restriction of the glaciers and ice-caps to less than 
their present extent and a corresponding rise in 
sea level. Geological evidence suggests that this 
rise may have been in the neighborhood of either 
2 or 3.5 m above present level. At that time, 
most of the atolls would have been submerged 
and have existed only as reefs awash with little 
or no dry land. With the gradual fall of sea level 
that started perhaps 3,500 years ago, platforms 
of reef limestone were exposed to erosional pro- 
cesses and to the deposition of the loose sedi- 
ments produced by this erosion. The resulting land 
habitats were a mosaic of area of sand and gravel 
bars, beaches, dunes, and flats, as well as reef 
breccia platforms variously eroded by action of 
rainwater. The material of this diverse substra- 
tum was limestone, pure except for possible minute 
amounts of drifted pumice. 

Weathering of this substratum by the solvent 
action of rainwater commenced, of course, imme- 
diately when it was exposed above the waves. This 
process was augmented by the development of 
vegetation even in its earliest stages, when very 
soon after coral limestone is exposed to air it is 
colonized by microscopic blue-green algae. Some 

forms that grow on solid surfaces have the ability 
to bore into the limestone. Others inhabit the 
interstices in the surface layers of sand-textured 
deposits, causing the particles to cohere and form 
a crust that is friable when dry and gelatinous 
when moist. The boring types speed the weather- 
ing of the limestone by breaking down the outer 
layers. Both types are suspected of contributing 
fixed nitrogen to the resulting soil, though this 
has not yet been demonstrated for the species 
involved. Certainly some humus is added as the 
plants die and decompose. 

The soil that forms in such situations is at best 
a poor one, fit only to support plants of the most 
absolute pioneer types. With the exception of 
calcium, the essential elements for plant nutrition 
are either scarce or chemically almost unavailable. 
Especially characteristic of these soils are high 
pH and deficiencies of nitrogen and iron, as well 
as of certain trace elements such as manganese and 
zinc. Phosphorus is initially low but may be 
increased very early by excrement from sea-and 
shore-birds. Nitrogen may be added in the 
same way. Extremely high salinity makes these 
soils even more unsuitable for most plants, as 
does the almost complete absence of organic 

As noted above, the development of microscop- 
ic blue-green algal vegetation, indicated by a 
darkening of the white or pinkish limestone, 
may tend to ameliorate the deficiency of organic 
matter and nitrogen. Even so, the number of 
plants fitted to colonize such a habitat is at best 
very low. This number is further restricted by 
the difficulties of dispersal over wide expanses of 
sea. The natural dispersal agents are practically 
limited to water, wind, and sea-birds. 

Almost the only members of the available floras 
that are adapted to colonize such essentially 
strand habitats are strand species, found every- 
where along the shores of islands and continents. 
These species are mostly adapted to dispersal 
by water, through various floating mechanisms, 
and by birds, through sticky or burr-like fruits 
that adhere to feathers or feet or through fleshy 
fruits that may be eaten and carried for some 
distance in birds' digestive tracts. A few species, 
such as some grasses, with very small fruits, and 



some plants with winged fruits, may be carried 
by winds, especially typhoon or hurricane winds. 

Recorded observations on the original compo- 
sition of the vascular vegetation on new coral 
atoll habitats are few. Pemphis acidula seedlings 
are known to become established in tiny pockets 
of sand caught on otherwise bare limestone 
rock. On coral sand and gravel banks, bars, 
or flats Portulaca lutea, Lepturus repens, Boer- 
havia spp., Guettarda speciosa, Tournefortia argen- 
tea, Scaevola sericea, Suriana maritima, Heliotro- 
pium anomalum, Jpomoea pes-caprae, Vigtia man- 
na, Pandanus tectorius, Cocos nucifera, Barringto- 
nia asiatica, and Thuarea involuta seedlings have 
been observed in absolutely unprotected appa- 
rently unmodified habitats. Of these, Cocos, 
Ipomoea, and Barringtonia have been seen in 
such circumstances only very rarely. Certain 
other species, though not observed as primary 
colonists on new areas, undoubtedly are able 
to fulfill this function. 

The simplest types of atoll vegetation are es- 
sentially aggregations of these pioneer species. 
In the driest atolls as well as in a few very remote 
ones that few species have reached, succession 
seems not to have progressed much beyond this 
stage where the original composition still charac- 
terizes the vegetation; the actual composition 
and local variations may result from chance or 
from local differences in substratum, position in 
relation to prevailing wind, distance from sea, 
or altitude. Characteristic types on such atolls 
are Scaevola scrub, scrub forest of Tournefortia, 
Guettarda and Scaevola; Lepturus grassland or 
Lepturus-Tournefortia savanna; mosaics of thicket 
and grassland; or dwarf scrub of Sida fallax or 
of Sida with Heliotropium. It is uncertain whether 
the Pisonia grandis forest, the Pisonia-Cordia 
forests, and the Pandanus forests in some of these 
extreme habitats represent pioneer stages or 
have developed after some slight alteration of 
the habitat and resulting succession. Mention 
should be made here of extreme halophytic types, 
such as mats of Sesuvium portulacastrum on 
lagoon margins and in saline depressions, stands 
of Pemphis where the roots are flooded by sea 
water at high tide, and scrub of Scaevola or 
Suriana bathed continuously by salt spray. 

The establishment of vegetation of any kind 
tends to bring about change and amelioration of 
the habitat in a number of ways. The simplest 
of these, of course, is shading. Even a slight 
protection from the heat of the sun, from the 
drying effect of the wind, and from salt spray may 
make possible the establishment of additional 

species or create more favorable conditions for 
some already present. Addition of humus to the 
soil tends to lower its alkalinity and to increase 
its water-holding and base exchange capacities. 
Phosphorus and nitrogen are increased by accu- 
mulation of bird excrement which, however, 
tends to be washed down through the porous soil. 
Nitrogen may also be contributed by the activity 
of bacteria on the roots of Vigna. of Azotohacter, 
and probably of blue-green algae. Nostoc is at 
times abundant on semi-shaded soil. A further 
effect may be the concentration of minor nutrient 
elements in humus. Rain tends to reduce salinity 
as well as to leach out nutrients, and to build 
up a body of fresh ground water at a slight depth 
in the soil. 

These processes are, of course, continuous and 
simultaneous, and usually make possible an 
increase in the flora and in the complexity of the 
vegetation. In submesic atolls, those with less 
than 2.5 m of precipitation, a mixed forest 
results which, in the Northern Marshalls, for 
example, may have all or any combination of 
about 16 species of trees and shrubs: Guettarda 
speciosa, Tournefortia argentea, Tenninalia samo- 
ensis, Morinda citrifolia, Pandanus tectorius, 
Pisonia grandis, Soulamea amara, Cordia subcor- 
data, Pemphis acidula, Ochrosia oppositifolia, 
Allophylus timorensis, Hernandia sonora, fntsia 
bijuga, Pipturus argentea, Scaevola sericea, and 
Suriana maritima. These may occur in any 
arrangement or combination with chance, dis- 
tance from the sea, and wetness of climate the 
principal factors involved. Such forests tend to 
be more luxuriant with greater rainfall. Substra- 
tum may be a factor in the pattern, though the 
solution is not clear except in a few cases such as 
where a pure or almost pure stand of Pemphis 
is found on bare rock. Many of the species listed 
may be dependent on shade for the establishment 
of their seedlings. 

In this mixed forest, ferns appear and mosses 
become more prominent. The ferns are principally 
terrestrial, and very few species occur. Certain 
successional phenomena may be observed, as the 
gradual dropping out of Tournefortia from the 
more dense forests, the assuming of dominance, 
in places, by single species, such as Pisonia, 
Ochrosia, Cordia, Intsia, or Pandanus. Forests 
made up exclusively of Pisonia arc especially 
frequent and occur under a wide range of mois- 
ture conditions. Almost no other species are 
found established under a thick stand of Pisonia. 
The factors involved in these changes are not 
usually obvious, and the matter needs much more 



careful study. With fairly abundant moisture, 
Ochrosia seems able to succeed any of the other 
species, even Pisonia, once it gains a foothold. 
Under Ochrosia its own seedlings usually form 
a veritable carpet, growing not more than a few 
cm high unless a break occurs in the dense cano- 
py, when they shoot up rapidly to fill it. 

It is of interest that the most stable, or most 
nearly "climax" types here seem to be composed 
of single species. 

In these submesic atolls, also, man has been 
able to establish an apparently permanent foot- 
hold. He has brought great alteration to much of 
the vegetation within his influence. Native 
forests have been cleared and replaced by coconut 
groves and plantations, and locally by mixed 
coconut and breadfruit, or by Calophyllum. In 
wet places he has planted taro, or rarely, man- 
groves. He has excavated pits down to ground 
water for raising taro and other crops. Later 
he has abandoned most of these in the submesic 
atolls. Increasing wetness of climate is reflected 
strikingly in greater luxuriance of coconut plan- 
tations and greater abundance of breadfruit trees. 

On the wetter atolls, the limitation on the de- 
velopment of vegetation seems to be much more 
the lack of available flora than the severity of the 
environment. The far greater richness of the 
vegetation of atolls in the western Carolines and 
especially in and near the Malaysian Archipelago, 
where a large flora is available, seems to support 
this idea. 

It is not clear whether the much greater pro- 
minence of mangroves in wet atolls is a function 
of climate or of their proximity to the rich 
mangrove swamps of Malaysia. 

Succession to a truly mesophytic vegetation, 
even to what might be termed rain forest, is in 
these wet atolls unimpeded either by dryness 
or by salinity. 

Studies of the vegetation of reasonably undis- 
turbed atolls of this character are as yet lacking. 
Those that have been accessible for investigation 
have been profoundly altered by man. The 
anthropogenic vegetation is correspondingly 
richer there than in drier atolls. In fact it is 
difficult at times to tell what species have been 
brought by man. The most striking major 
aspects are solid, dense forests of giant bread- 
fruit trees and extensive taro-pits or marsh gar- 
dens in which many food plants in addition to 
taro are grown. The weedy vegetation and the 
undergrowth in coconut plantations are both 
more luxuriant and richer in species. 

Generally speaking, after the earlier stages of 
colonization and development of vegetation have 
been passed, the differences in atoll vegetation 
in the Pacific may be interpreted in terms of 
three major gradients that from dry and seasonal 
to wet climates, that from saline to fresh sites, 
and that from the east, with a very small available 
flora, to the west and southwest with the rich 
Malaysian and Melanesian floras. 

All three of these gradients, as expressed, 
exhibit increasing richness of composition of 
vegetation. Toward their extreme ends of 
dryness, salinity, and remoteness, the tendency 
to complete dominance by single species is marked. 
Such types dominated by single species, are rare 
in mesic atolls excepting the forests of Pisonia 

The elTects of decreasing salinity are reflected 
in the increased luxuriance and richer composi- 
tion of vegetation in the interior of fairly wide 
islets, and, very strikingly, by the great increase 
in number of species on even slightly elevated 
atolls over those lying at sea level. The increase 
on the latter type, however, may also be inter- 
preted as a result of greater age of the land surface 
in raised atolls. 

From dry to wet atolls, both luxuriance and 
number of species increase in a striking fashion, 
whether we are considering only native vegetation 
and flora or including that established by man. 
The two driest atolls of the northern Marshalls 
have vascular floras of nine species each, while 
some of the wet southern atolls of the same group 
have as many as a hundred or more species. 

Accurate figures to show the effect of remote- 
ness are not available for atolls at the ends of the 
gradient; but it is possible that Clipperton 
Atoll had no native plants, and only two are 
recorded from Ducie Atoll. Vostok and John- 
ston Islands, not so far to the east, but very 
remote, had respectively two and three species 
of native vascular plants. However, the floras 
of other atolls in southeastern and central Poly- 
nesia are more diverse than these, but nothing 
to compare with the large numbers on various 
East Indian Atolls, or even the Caroline ones. 
Accurate floristic figures are much to be desired 
for atolls in the Malaysian area. 

It is thought that in very few atolls has vegeta- 
tion developed or succession reached its maxi- 
mum possible development under present climatic 
conditions, as the soils are very immature and 
oversea migration of plants is sporadic and 


uncertain. Where atolls have been left undis- owing to the disturbances caused by hurricanes 

turbed, though, the climatic conditions have and by man's activities, most atoll vegetation 

tended to arrest development at different points, may be regarded as being in a more or less active 

corresponding to degrees of wetness. However, dynamic status. 





Pacific Vegetation Project, Care of National Research Council, Washington, D.C., U.S.A. 

Grassland vegetation, a collective term for 
plant communities in which grasses and grasslike 
plants are the dominant growth form, is fairly 
widely distributed in the tropical Pacific Basin 
but in aggregate makes up only a small part of 
the vegetation. Natural, as compared with man- 
made, grassland covers an even smaller area. 
In some instances it is difficult to be certain of the 
history of a given area. 

The classification of grassland may be based 
on any of several features composition, physiog- 
nomy, structure, habitat, or economic considera- 
tions. Because no adequate study has been made 
of the region as a whole on any of these bases, and 
because the required information is not available 
on any of these features for all Pacific grasslands, 
it seems logical to base a tentative arrangement 
on such data as are available on all features. 
Many tropical grasslands, including some Pacific 
ones, contain scattered trees and are then gen- 
erally called "savanna." Every stage exists 
between grassland without trees through savanna 
and woodland to forest, depending on the spacing 
of the trees. A savanna may be arbitrarily defined 
as grassland with trees that are spaced, on the 
average, farther apart than twice the diameter 
of their crowns, or with trees covering less than 
20 per cent of the ground. The significance of 
trees in grassland varies, depending on the kinds 
of trees. In some cases they may represent 
invasion leading to re-establishment of forest; 
in others they may belong to species that are 
able to resist burning when well grown, but 
which fire will kill when young. Still other 
savanna trees, usually small ones, belong to 
species that are found nowhere else than scat- 
tered in grassland. Considerable work has been 
done on savannas in other parts of the tropics 
but rather little on those of the Pacific. Here they 
will be treated mostly as variants of grassland 
types and discussed individually in the above 

The Pacific grasslands may be conveniently 
separated into those of high altitudes and those 
of medium to low altitudes. Fixed altitude limits 
may not easily be assigned to these two categories, 
but the high-altitude types either are mostly 
natural or are moderate expansions of natural 

grassland, while many, if not most, of the middle 
and low altitude types seem to result from or to 
have been vastly extended by the activities of man. 

Montane grasslands are extensively developed 
in the Pacific Basin principally in the Andes 
where they occupy the area between timber line 
and perpetual snow. Timber line ranges from 
about 10,500 to 1 1 ,000 feet in Colombia to nearly 
13,000 feet in Bolivia, and there is a correspond- 
ing variation in snow line from perhaps 15,000 to 
18,000 feet. The Andean montane grassland is 
a bunch-grass or tussock formation with Stipa, 
Calamagrostis, Poa, Agrostis, and some other 
genera as the principal grasses and with abundant 
cushion plants and a rich herb and shrub flora. 
This grassland belt stretches from Patagonia to 
northernmost Colombia and, in poorly developed 
form, to Costa Rica. It is continuous in the 
south, but is broken into isolated patches in the 
lower northern Andes. Still, even in Colombia, 
the aggregate area is large (3, 4, 5). 

In addition to much local variation, correlated 
with altitude, topography, hydrology, exposure, 
and human interference, there is a large-scale 
regional variation from north to south, related 
to climate. This variation expresses itself in three 
major grassland regions or formations, designated 
conveniently by the local names "pramo," 
"jalca," and "puna." They occur in that order 
from north to south and are characterized by 
increasing aridity southward. 

The paramo is the most remarkable of the three, 
marked by features that might well lead to the 
argument that it is not a grassland at all. The 
most striking feature is the remarkable composite 
genus Espeletia. The grotesque gray rosettes of 
this group, called "frailejones" by the local people, 
are an almost constant feature of the landscape 
above timber line from the Venezuelan Andes to 
the Paramo del Angel in northern Ecuador, where 
they may be seen in great abundance, but where 
their occurrence stops abruptly. South of the 
Rio de los Charcos they are completely unknown. 
Although usually forming only a small part of 
the cover, Espelatia gives a unique aspect to the 
landscape that amply warrants separation of the 
pdramo from the jalca to the south. In addition, 
there is a very considerable flora of herbaceous 



and shrubby plants with striking local endemism 
which sets off this wetter northern paramo region. 
Giant bromeliads (Puya spp.), Rumex tolimensis, 
Lupinus alopecuroides, Aragoa spp., Blechnum 
arboreum, and other strange plants add to the 
unreality of this landscape. 

On level or more gently sloping ground, the 
soil is usually black and highly organic. Rainfall 
is high, fog is frequent, and mosses, ferns, and 
other hygrophytes are in places very abundant. 
The arrangement of the mountains somewhat 
parallel to the direction of the trade winds lessens 
possible rain-shadow effects. 

The paramo vegetation thins out upward to 
the level of perpetual snow. The upper part, 
or super-paramo is more strictly grassland, though 
sparse. Downward, the shrubby sub-paramo 
vegetation gradually replaces the grassland in a 
transition to the "sotobosque" or "elfin forest" 
and the mossy forest. 

Disturbance by man in the region just below 
the paramo has had the interesting effect of caus- 
ing at least some elements of the paramo vegeta- 
tion to spread downward in places, producing, 
what might be called quasi-paramo, very similar 
to the paramo in aspect down to elevations as low 
as 3,000 m. But many paramo species are lacking, 
and there are many introduced weedy species, 
including grasses, not normally found in the 
paramo. Active human utilization of the pdramo 
does not extend much above 3,500 m. 

From the Charcos River in northern Ecuador 
to north-central Peru, the jalca (often also 
called paramo or even "wet puna") occupies a 
broadening belt, similarly placed with regard to 
the forest belts below and perpetual snow above. 
It is a grassland of bunch-grass or tussock species, 
with, again, many non-graminoid herbs and 
shrubs, but lacking the Espeletia and thus present- 
ing a far more conventional mountain meadow 
appearance. This is varied, however, by such 
oddities as Puya, Azorella, and Distichium. 

The jalca, though mostly somewhat drier than 
the paramo, is still a mesophytic grassland, and 
the ground cover is complete. Fog is still an 
important factor in the climate. Human occupa- 
tion extends locally to 4,000 m elevation. A 
limiting factor seems to be the cold, resulting 
from the high degree of cloudiness. 

Southward, where the climate becomes signi- 
ficantly drier, the grass clumps become more 
discrete; Stipa becomes the dominant genus; 
and bare ground becomes apparent, at least locally 
between the clumps. This is the beginning of the 
"puna" or high altitude "steppe." 

Here timber line, on the east slopes, is higher. 
Woodland of Polylepis reaches 4,200 m. The 
associated herbs and shrubs in the grassland are 
mostly more xeromorphic. Such genera as Te- 
traglochin, Pycnophyllum, Lepidophyllum, and 
Opuntia are found. Large woody species of the 
cushion plant, Azorella, are common and locally, 
along with the shrub Lepidophyllum, provide fuel 
for the inhabitants. 

Here human activity normally extends to 5,000 
m, and a road crosses a pass in Bolivia at over 
6,100 m. This increase in the altitude limits of 
effective human occupation seems due to the 
higher insolation and lower degree of cloudiness. 

Enormous areas of puna occupy the broad part 
of the high Andes, the "altiplano" of southern 
Peru, Bolivia, and Chile. On the east slope it 
merges with the "ceja de la montana" or elfin 
forest of the Yungas, on the west into the desert 
western slopes of the Andes in Peru and Atacama, 
Chile. Southward it extends, at lower and lower 
altitudes, to the south temperate zone and even, 
in modified form, to Patagonia. 

Elsewhere in the tropical Pacific the only equi- 
valent of Andean high-altitude grassland is a 
weakly developed belt in Hawaii. This is charac- 
terized by sparse stands of densely tufted species, 
up to 1 m tall, mainly Trisetum glomeratum, 
Agrostis sandwicensis, and Air a nubigena, at timber 
line on the high volcanoes above 2,600 m. The 
belt merges downward with the upper forest and 
upward into bare lava, cinder, and clinker slopes. 

In the saddle between Mauna Loa and Mauna 
Kea on Hawaii, similar grassland is dominated 
by tufted perennial species of Eragrostis. Exces- 
sive grazing by sheep and wild goats has damaged 
and altered these grasslands. 

Mention may also be made here of extensive 
pastures existing at middle altitudes on the north- 
ern part of the island of Hawaii. These are 
artificial and are made up of numerous species of 
forage grasses deliberately introduced for the 
raising of beef cattle. 

Alpine grassland, in the mountains of Papua, 
dominated by Aulacolepis, Poa, and Danthonia, 
are mentioned by Brass (2, p. 177), but no des- 
cription has been found. 

At low and middle altitudes in the Pacific 
tropics, there are of course large expanses of such 
cultivated grasslands as sugar cane and rice fields. 
Various minor grassy communities, such as 
marshes, old fields, salt-grass flats and other strand 
types, and pastures are of reasonably general 
distribution. Worthy of special mention are the 
Lepturus repens grassland of coral atolls and other 



strand situations and the reed marshes composed 
of Phragmites karka. Stands of Lepturus are 
widespread in small patches on dry coral sands 
and cover substantial areas on Christmas Island 
and other dry atolls of the Central Pacific and on 
Pokak Atoll, the driest of the Marshalls, but are 
strangely missing in Hawaii. 

Lepturus repens, in spite of its name, is a bunch 
grass, and occurs in discrete rather small tufts of 
fine but hard culms, and narrow but harsh leaves. 
Frequently these tufts will send out long wiry 
runners which root at the nodes and send up 
smaller tufts. Eventually a loose mat may be 
formed. Usually one or more of several broad- 
leafed herbs and dwarf shrubs occur mixed with 
the grass, in varying proportions. These are 
Portulaca lutea, Boerhavia sp., Sida fallax, and 
Heliotropium anomalum. The last is lacking in 
the Marshalls but found in Wake and the Line 
islands. Scattered low, rounded trees of Tourne- 
fortia argentea may be found, making a savanna 
of very characteristic appearance. 

Reed marshes are widely distributed on higher 
islands, usually, but by no means always, 
near sea level just back of the coasts. Fairly 
extensive marshes are found on Saipan, Guam, 
and Truk islands. They are usually almost pure 
stands of the large tropical reed, Phragmites karka. 
It forms hollow canes about 1 to 1.5 cm thick, 
with broad leaves above, and with panicles 
bronze-colored at flowering. Pandanus trees and 
Hibiscus tiliaceus may occur scattered through 
these marshes to form savannas. 

Much more extensive are several upland types 
of grassland or savanna. Most widely distributed 
of these, though possibly not largest in area, is 
that dominated by the genus Miscanthus, prin- 
cipally by Miscanthus floridulus, commonly called 
"sword-grass" because of the scabrous cutting 
edges of its leaves. 

In its simplest form, Miscanthus grassland is a 
pure stand of sword-grass, crowded large tufts of 
reed-like culms with harsh leaves, up to 3 and 
even 4 m tall. It is found thus on relatively 
fresh ash slopes, as on the volcanoes of the 
northern Marianas. Reasonably pure stands are 
also widespread in denuded areas on the older 
volcanic islands of southeastern Polynesia, es- 
pecially Mangareva. Generally, however, there 
is a varied accompanying flora of sedges, other 
grasses, and several broad-leafed species of herbs 
and shrubs. Most of these grow on erosion scars 
and other bare spots, which are an almost con- 
stant feature of the habitat of Miscanthus. In 
Guam, the Miscanthus grassland forms a mosaic 


with a low, soft grassland composed mainly of 
Dimeria chloridiformis. In this mosaic, Dimeria 
occupies principally the more level or gently 
sloping situations, fine clay soil that dries out 
fairly well in dry weather. The steeper, more 
rocky sites, as well as those with more permanent 
moisture, are covered predominantly by Miscan- 

Fire is a frequent feature of Miscanthus grass- 
land and the associated Dimeria. In burned areas, 
an aggregation of weeds appears, to be gradually 
crowded out by the grasses. If erosion starts, 
an erosion scar community of ferns, small shrubs, 
and various herbs tends to establish itself on 
the bare soil. As time goes on, this community 
is replaced by the grasses. Most of the plants 
peculiar to these grasslands are members of this 
erosion scar community. Members of the genera 
Myrtella, Melastoma^ Geniostoma^ Hedyotis, 77- 
monius, Dianella, Euphorbia, Phyllanthus, Glochi- 
dion, Machaerina, Rhynchospora, Fimbristylis, 
Cantharospermum, Chrysopogon, and Ischaemum, 
as well as the ferns Gleichenia, Sphenomeris^ 
Lindsaya, and Cheilanthes, plus Lycopodium, 
make up the community. They are mostly either 
very wide-ranging species or very restricted 

The bottoms of permanently or usually wet 
ravines in the mosaic are occupied by brakes of 
reeds (Phragmites karka) which show up as a 
deeper green or, at flowering time, a rich bronze 
against the pale green or buff of the Miscanthus. 
These reeds are very tall, often filling ravines up to 
the level of the grass tops on the terraces along 
the ravine margins. Only the difference in color 
betrays the existence of the ravine to the casual 

In this mosaic, seedlings of Casuarina equiseti- 
folia tend to appear; and an area not burned for 
a few years may assume the character of a savan- 
na, or even of a sparse forest. Pandanus also form 
limited savanna. 

Westward in the Pacific, other grasslands tend 
to replace Miscanthus as the important types. 
In Yap and Palau, savannas of ferns, Nepenthes, 
and sedges with some grasses and many secondary 
herbs and scattered trees and shrubs of various 
kinds occupy the places where Miscanthus would 
be expected. 

In the Philippines and northward, Miscanthus 
floridulus tends to be replaced in certain situations 
by Miscanthus sinensis. The exact environmental 
relations of these two species are not always 
clear, nor is it easy to distinguish them under 
some conditions. In the Ryukyus where Miscan- 



thus may be unusually abundant on limestone, 
in contrast to its behavior farther east, the species 
involved may often be Miscanthus sinensis though 
this has not been determined with certainty. No 
information is available on the Miscanthus (?) 
grasslands found in western Papua between 1,000 
and 2,000 meters of elevation and mentioned 
by Brass (2, p. 176). 

In places in the Philippines, Caroline Islands, 
and probably elsewhere, Saccharwn spontaneum, 
a large bunch grass with the aspect of Miscanthus, 
may form considerable stands. Not too much is 
known of the distribution of this type of grassland. 
Since the war, Saccharum spontaneum has ap- 
peared in Saipan and spread with great vigor. 
It is suspected that the plants involved may really 
be seedlings from cultivated sugar cane. True 
Saccharum spontaneum^ course, may well have 
been introduced accidentally with war supplies, 
or perhaps after the war. The species seems na- 
tive and not excessively abundant in the high 
Caroline Islands from Kusaie to Palau, where it 
is frequently mistaken for Miscanthus. The 
latter is known with certainty in the Caroline 
Islands only from Ponape. 

In the Fly River area in New Guinea and west- 
ward are large areas of savanna (2), dominated 
by Ophiurus exaltatus, a coarse grass reaching 2 
m in height, but also characterized by Themcda 
triandria, Imperata cylindrica^ and Ischacmum sp., 
and scattered trees of Eucalyptus, Melaleuca, 
and Tristania. Tristania dominates or shares low, 
wet places with Eriocaulon and many other herbs. 
In places, Ophiurus, Imperata, and Sorghum 
nitidum form a grassland on well-drained soils. 
Elsewhere, Germainia ca pi tat a forms savannas 
with Banksia. Low, ill-drained soils have a 
ground cover of sedges with numerous other 
flowering herbs and small trees of Melaleuca and 
Banksia. These savannas fade imperceptibly into 
"savanna forests" or woodlands. Brass considers 
these savannas to be natural and to be extensions 
into New Guinea of the "open forests" of Aus- 

From Guadalcanal to the Philippines, and 
undoubtedly westward in Indonesia in such 
islands as Celebes, Timor, and the lesser Sunda 
group, occurs a rather coarse grassland domi- 
nated by Themeda triandria, the "kunai" grass of 
New Guinea. In Guadalcanal, Florida, and 
probably Buka in the Solomons, an almost pure 
stand of Themeda, with wet places dominated by 
Phragmites karka (9, pp. 90-91) occurs on the 
coastal plain and runs up onto low mountain 
ridges several miles inland, where it changes to 

savanna and gradually to forest upward. Theme- 
da triandria is a stiffish grass 1 to 2 meters in 
height, with drooping flower clusters. 

In the Philippines, the Themeda type of grass- 
land is the matrix of many of the pine savannas 
of Benguet, Luzon. Themeda triandria, Themeda 
gigantea, Chrysopogon aciculatum, species of 
Ischaemum, Panicum, Andropogon, and Fimbris- 
tylis, as well as various other grasses, sedges, and 
broad-leafed plants make up this grass cover. 
Merrill (8) mentions large areas of Themeda- 
dominated grassland throughout the Archipelago 
and lists a large number of species as common in 
open grasslands. There are obviously many 
variants of this type as well as of the Imperata 
grassland to be described next. 

Although Themcda and its associates are main- 
tained by burning, if the burning is carried to an 
extreme, this type of grassland may give way to 
Imperata cylindrica and associated species. 

Imperata grassland covers vast acreages in the 
New Guinea-Indonesia-Southeast Asia region, 
and is usually, if not always, associated with man. 
Its altitudinal range is enormous from sea level 
to as much as 2,500 m (in New Guinea, according 
to 6). Imperata is the "lalang" of Malaya, the 
"alang-alang" of Indo China, the "cogon" of the 
Philippines. It may be thought of as the true 
"climax" of the man -shifting-cultivation-burning 
complex in this part of the world. Imperata, with 
the tough matted wiry buried rhizomes, is about 
as near as nature comes to producing a really 
fire-proof plant. When the shoots are burned 
off, the rhizomes remain uninjured and immediate- 
ly respond by sending up flowering culms, fol- 
lowed by a new crop of leafy shoots. 

Though Imperata is a harsh, unpleasant grass 
in texture, it is not large. It forms a tough hard 
sod, is not bunchy, and seldom exceeds 1 meter 
in height. Its bright green color gives a deceptive 
appearance of succulence, but animals do not 
care much for it. It is commonly associated with 
a rather characteristic lot of widespread plants 
belonging to such genera as Melastoma, Pteridium, 
Lygodium, Eurya, Gleichenia, Rhynchospora, 
Fimbristylis, and Andropogon. 

Although this type of grassland is easily invaded 
by woody species, such as pine, such invasion 
seldom leads to anything, as the invaders are 
generally susceptible to fire. And fire, the factor 
that brings about dominance by Imperata, is 
seldom long absent where Imperata has achieved 
such dominance. 

Much study has been devoted to the origins of 
the grasslands and savannas of tropical America 



and Africa. A paper by Beard ( 1 ) provides a 
convincing set of generalizations about at least 
the tropical American savannas. The grasslands 
of the western tropical Pacific have received little 
attention, as it has generally been assumed that 
they arc of secondary origin, due to man's activ- 
ities. At least one student (9) has vigorously 
contested this view as applied to the grassy areas 
of Guadalcanal, withal not too convincingly. 
His conclusions are largely based on low rainfall, 
as shown by a very short period of record; on 
absence of evidence of burning at the present 
time; on the existence of strips of forest along 
streams; on the existence of two peculiar animals, 
an endemic subspecies of button quail an obli- 
gate grassland bird and a fish adapted to inter- 
mittent streams; and especially on a curious 
argument that, because the climate cannot support 
rain forest the grassland must be "climatic." No 
mention is made of the possibility of other types 
of tropical forest. From the data he presents, 
there is no doubt whatever that these grasslands 
depend for their existence on the rain shadow 
cast by the high Guadalcanal mountains. Whether 
they are climatic in origin is another matter. 
They may very well be so, but it is not established 
to complete satisfaction in his paper that man in 
earlier times did not have a hand in their origin 
or spread. The fact that he describes them as an 
almost pure stand of Themeda triandria and men- 
tions almost no accompanying flora makes their 
natural origin at least open to question. 

Similar views concerning the Miscanthus grass- 
lands of Fiji and Guam have been expressed oral- 
ly by a number of casual observers. In the case of 
Fiji, these views were based on the low rainfall, 
and in that of Guam on edaphic considerations. 

Some study has been devoted to the origin of 
the Guam grasslands. Here, on historical evi- 
dence, there was much more forest even within 
historical times. The introduction of deer and 
of the custom of driving them by fire caused a 
great reduction of this forest during Spanish times. 
Many of the plants characteristic of the present 
grassland were clearly introduced from America 
and elsewhere during the Spanish period. These 
facts favor the idea that this grassland is of 
secondary nature. However, as in tropical 
America, there are some native, even endemic 
species, restricted to the grasslands. Such are 
Phyllanthus saffordii, Ischaemum longisetum, 
Machaerina aromatica, Hedyotis grandiflora, Wik- 
stroemia elliptica, and Spaihoglottis micronesiaca. 
Glochidion marianum is found on both Guam and 
Ponape; Myrtella benningseniana, Timonius albus, 

Hedyotis fruticulosa on Guam and Yap; and 
Dimeria chloridiformis on Guam and Palau. 

In addition there are certain widespread plants 
of similar habitats that are not obviously intro- 
duced, such as Miscanthus floridulus and Dianella 
ensifolia (both known as fossils from Pagan, 
northern Marianas), Curculigo orchioides, Fim- 
bristylis tristachya, Fimbristylis annua, Melasto- 
ma malabathricum, Lindsaya ensifolia, Cheilanthes 
tenuifolia, and Lygodium seandens. 

This assemblage of plants could not likely 
have developed or lived in forest, as it is made up 
of shade-intolerant species which do not now 
inhabit nearby forest. The obvious possible 
habitats were considered. Fresh ash slopes, 
such as those covered by Miscanthus in the 
northern Marianas, were ruled out, as such have 
apparently never existed in Guam. Open ridge 
crests do not seem likely, as in a climate such as 
that of Guam and at such low altitudes the ridges 
are likely to have been wooded. Also many of 
the species concerned do not elsewhere inhabit 
such ridge crests. Landslide and erosion scars 
would be a possibility for some of the species, if 
there ever had been an abundance of landslides, 
but this does not seem a sufficient possibility, in 
the absence of humans and grazing animals to 
cause large numbers of landslides and aggravated 
erosion. The late Josaiah Bridge suggested a 
correlation with bauxite deposits. This would 
be a possibility, as many of the same plants are 
associated with bauxite elsewhere. However, 
extensive deposits of bauxite are lacking in the 
areas concerned. In any event, even with bauxite, 
open habitats would have been required. 

In attacking the same problem in the tropical 
American savannas, Beard (I) noted a constant 
association of what he regarded as natural savan- 
na with areas of mature topography flat or gently 
undulating country, on plains, terraces or plat- 
eaus, having soil with impeded drainage, generally 
a permeable layer lying on an impermeable one. 

It seemed pertinent to look for such an asso- 
ciation of savanna and topography on Guam, 
past or present. When asked about this, J.I. 
Tracey suggested that certain flat-topped erosion 
remnants scattered over southern Guam might 
represent the remains of an ancient flat erosion 
surface. These "mesitas" present precisely the 
flat surface and permeable surface layer overlying 
a clay subsoil that are required by Beard's theory. 
The general geological reconstruction of southern 
Guam indicates that this flat surface may have 
been in existence for a long period, possibly since 
some time in the Miocene. If this is so, a proper 


habitat would have been available for the develop- 
ment of the native part of the floristic assemblage 
now characteristic of the Guam grasslands. 

It is suggested that the history of these grass- 
lands is about as follows : 

1. The gradual appearance of grasslands and 
their flora, and their encroachment on the forest 
as the topography matured during the erosion 
cycle following the most recent emergence of 
most of the area from the middle Miocene Alifan 

2. The gradual restriction of these grasslands 
and encroachment of forest as the flat surface was 
dissected during elevation and rejuvenation. 

3. The persistence of grassland only in small 
patches, mostly on the "mesitas" or erosion rem- 
nants during the time of the Chamorro inhabi- 
tants of Guam. 

4. The rapid and extensive expansion of the 
grasslands onto the denuded eroded hills, with 
addition of numerous introduced species from 
the time of the Spanish occupation to the present 

Similarity in situation on other Micronesian 
islands, such as Palau and Yap, suggests that 
possibilities of a similar history might be inves- 
tigated when the necessary geological information 
is available. Until floristic, geologic, and soils 
data can be assembled for other Pacific grasslands, 
little more can be said about their origin beyond 
the statement that in most regions they have at 
least been greatly extended, if not actually brought 
into being, by human agencies. 

Merrill (7, 8) categorically considers the 
Philippine grasslands to be the result of human 

activity. The fact that some of the plants found 
in these grasslands are endemic to the Philippines 
suggests that this may be an extreme interpretation 
and needs more study. However, it is easy 
enough to see that the pine savannas of Luzon are 
increasing their area very conspicuously at the 
present time. Certainly it is safe to assume that at 
least most of the Philippine grasslands are secon- 

Descriptions of the New Guinea savannas (2) 
suggest that at least some of them may be natural, 
with the same characteristics of alternate extreme 
desiccation and waterlogging described by Beard 
for tropical America. These areas would amply 
repay further study. 


(1) Beard, J., 1953, Ecol. Monogr. 23: 149-215. 

(2) Brass, L., 1938, Jour. Arnold Arb., 19: 


(3) Cuatrecasas, J., 1934, Trab. Mus. Nac. 

Cicnc. Nat. Ser. Bot., 27: 1-144, 

(4) ..... ___ ., 1956, Suelos Ecuatoriales 1: 


(5) Fosberg, F.R., 1944, Jour. N. Y. Bot. Card. 

45: 226-234. 

(6) Lam, H.J., 1945, Sargentia 5: 153, 168. 

(7) Merrill, E.D., 1912, Phil. Jour. Sci. Bot. 7: 


(8) __ _ , 1923-1926, An enumeration of 

Philippine flowering Plants. 4 vols., 

(9) Pendleton, R.E., 1949, Ecol. Monogr. 19: 






Laboratory of Plant Ecology, Institute of Polytechnics, Osaka City University, Osaka, Japan. 


In the ordinary sequence of climatic zones on 
the earth, the moist climates of temperate and 
tropical regions are separated from each other by 
the intervening dry climate which prevails over 
middle latitudes. So that the two major forest 
areas on the continents, temperate and tropical, 
rarely come into contact in their geographical 
distribution. This geographical isolation has made 
plant ccologists overestimate the contrast between 
temperate and tropical forest vegetation in phy- 
siognomy, community structure and floristic 
diversity. Indeed the luxuriant growth of huge 
trees as well as the immense richness of plant 
species in tropical forests is a wonder to those who 
are acquainted with temperate vegetation, but 

nevertheless it is not right to lay too much stress 
upon this first impression. 

Along the western fringe of the Pacific Basin, 
a long chain of islands stretches from north to 
south along the continent, starting from Sakhalin 
through Japanese Archipelago, the Ryukyus and 
Formosa to the Philippines, where both summer 
and winter monsoon bring abundant rainfall 
throughout the year. It should be emphasized 
that this part of the western Pacific is perhaps the 
sole example in the world that represents the un- 
interrupted series of moist forest climates from 
the subarctic to the tropical zone. According to 
the researches of Japanese ecologists (1,2,3), no 
less than six forest zones are discriminated along 
the western coast of the Pacific. The parallel 
nomenclature of climatic and vegetation zones 
are as follows. 


0. Polar zone 

1. Arctic zone 

2. Subarctic zone 


month-degrees Barrens, snow and ice 

0-15 Tundra 

15-45 A 55 Evergreen conifer forest Abies, Picea, Larix, Betu- 

la, etc. 

Deciduous broad-leaf Fagus, Quercus, Acer, etc. 

Lucidophyllous or Cyclobalanopsis, Shiia, 
laurel forest Machllus, etc. 

,, Castanopsis, Lithocarpus, 

Cinnamomum, etc. 

Subtropical rain forest Ficus, Machilus, Lager- 
stroemia, and many other 

240 - Tropical rain forest Floristic composition 

extremely diverse. Genera 
of Dipterocarpaceae are 
most remarkable 

* Warmth index is a convenient geographical index of thermal climate proposed by Kira (1). It is given by summing 
the mean monthly temperature (t) above 5C or by I (/ - 5). Winter months in which / < 5C are excluded. 


3. Cool temperate zone 45 A 55 - 85 

4. Temperate zone 85 - 140 
41. Warm temperate zone 140-180 

5. Subtropical zone 180-240 

6. Tropical zone 



These systems also proved applicable to the 
altitudinal zones on high mountains of Formosa, 
continental China, Himalaya (4,5), and Malay- 
sia (2), with slight modifications. The zones 
presented here seem more in number than those 
in any other systems hitherto proposed. This is 
not strange because the differentiation of vege- 
tation zones along thermal gradient is expected 
to be most complete under moist climates. Where 
the subhumid or arid climate interferes with the 
normal sequence, the thermal zones may reduce 
in number or even be discontinued as is usually 
the case in other parts of the world, on which 
most other systems of zonal classification have 
been based. 

These zones are more or less physiognomical. 

They do not represent the discrete units of vege- 
tation, but are no more than the vague patterns 
joined together by gradual transition. As stressed 
by Imanishi since 1937 (6,7,8) and recently by 
Curtis (9 JO) and Whittaker (11J2), one zone 
gradually and continuously gives way to another, 
when the vegetation is analysed in terms of the 
distribution of each component species along an 
environmental gradient, e.g., thermal, moisture, 
etc. Fig. 1 illustrates an example of the distri- 
bution of some twenty conifer species in central 
Japan along the thermal gradient, including both 
horizontal and altitudinal. As seen in the figure, 
all distribution curves are bell-shaped and neither 
the ends nor the modes of curves tend to con- 
centrate into definite groups, although the area 

Warmth index (month -degrees) 

Fig. 1. Distribution of 24 conifer species along thermal gradient in the Central District, Japan. 1 :Pinus pumila. 
2 : Pinus koraiensis. 3 : Abies Mariesii. 4 : Abies Veitchii. 5 : Picea jezoensis var. hondoensis. 6 : Larix Kaempferi. 
1 : Tsuga diver sifolia. 8 : Picea bicolor. 9: Tax us cuspidal a. 10 : Abies homolepis. 11 : Pinus pentaphylla. 12 : Thuja 
Standishii. 13 : Chamaecyparis pisifera. 14 : Picea polita. 15 : Chamaecyparis obtusa. 16 : Thujopsis dolabrata. 17 : 
Tsuga Sieboldii. 18 : Sciadopitys verticillata. 19 : Abies firma. 20 : Cryptomeria japonica. 21 : Pinus Himekomatsu. 
22 : Pinus densiflora. 23 : Torreya nucifera. 24 : Pinus Thunbergii. 



surveyed includes four zones from arctic (alpine) 
to temperate. 

In the similar way, as we travel southwards 
from the subarctic conifer forest of Sakhalin and 
Hokkaido, the nature of forest community gradu- 
ally but steadily changes towards the tropical 
rain forest of the Philippines. No discontinuous, 
sudden change is found except where the flora is 
impoverished due to the small size of the islet as 
on the Tokara Group at the southern end of 
Japan, or where the thermal gradient is extra- 
ordinary sharp as in the narrow straight between 
Formosa and Botel Tobago. Thus we are inclined 
to consider that the aforementioned difference 
between temperate and tropical forest is nothing 
but the contrast between both ends of a con- 
tinuous series. 

As an instance, the floristic diversity of the 
tropical rain forest, so often spoken of in ecolog- 
ical literatures, will be considered from this view- 
point. In recent years, plant and animal ecolog- 
ists have recognized that the number of species 
and the number of individuals per species in a 
community are subject to certain statistical regu- 
larities. Let us now adopt Fisher's law of 
logarithmic series ( 13) . Denoting by s the number 
of species which is represented by n individuals 
in the sample randomly collected from a certain 
community, Fisher's law is given by 

s = a 

x n 

where x (0 < x < 1) is the value that represents 
the completeness of sample and the more com- 
plete is the sample the nearer it approaches unity. 
a, on the other hand, is independent to sample 
size and means the number of species represented 
by a single individual when the community is 
completely sampled. The latter value or the 
index of diversity offers an exact measure of esti- 
mating the floristic diversity of a community. 

As shown in Figs. 2 and 3, the law is fairly well 
satisfied by forest samples of utterly different 
nature and from different climatic zones. Es- 
pecially noteworthy is the fact that a single domi- 
nant forest in the temperate zone (Fig. 2) and a 
tropical rain forest that contains nearly 100 tree 
species per hectare (Fig. 3) equally well fit the 
law. Apparent correlation is found when the 
values of a obtained from different climatic zones 
are plotted against thermal gradient (Fig. 4). These 
evidences suggest the continuous decline of 
floristic diversity from tropics to high latitudes 
and do not support the view that the tropical forest 
is essentially different in its character from the 
temperate forest. 


In his monograph on the tropical rain forest, 
P.W. Richards (4) has recently stated that the 
tropical rain forest is not a highly specialized plant 
community as usually believed, but represents 
the most original type from which more impov- 
erished vegetation of temperate regions have 
been derived through evolutional history. With 
regard to floristic diversity, he found that the 
diversity was most pronounced in the mixed forest 
on mesic sites, and that, even under the same 
rain forest climate, forests on perhumid or sub- 
humid sites tended to be poorer in tree species 
and to have a few more or less apparent domi- 
nants. A similar principle might also be applied 
to the changes due to the fall in temperature. 


In view of the continuous concept of the world's 
vegetation as stated above, the study of tropical 
forests, especially of the mixed rain forests, is 
doubtlessly one of the most important subjects 
for plant ecologists, as it may justly be called the 
starting point of all kinds of vegetation study. 
Owing to the unfavorable climate and inaccessi- 
bility from civillized areas, however, researches 
on tropical forest ecology in the past were mostly 
confined to community floristics and morphology. 
Among the problems left for future study, we are 
especially interested in the productivity of forest 
ecosystems under rain forest climate. 

As early as in 1930, Vageler (15) estimated the 
annual organic matter production by tropical 
rain forest to be 100-200 tons per hectare. There 
is little doubt about the widespread conception 
that tropical rain forest is by far the most pro- 
ductive of all terrestrial ecosystems. Vast accumu- 
lations or standing crop of living plant body on 
unit forest area is mainly responsible for the con- 
ception. But the amount of standing crop is not 
always proportional to the true productivity or 
the velocity of organic matter production, usually 
given by the amount produced per year and per 
unit area. As a considerable portion of annual 
production is accumulated as wood year after 
year, and leaves as well as small branches and 
roots are continuously shed off day by day, the 
productivity of a forest can not be estimated from 
the standing crop alone. 

The difficulty in determining the true productiv- 
ity caused considerable variation in the estimates 
by different authors even for temperate forests. 
Filzer (16), for instance, estimated the dry matter 
production of German forests to be ca. 4.6 t/ha/ 
year. This figure was largely common, he stated, 

to pine, spruce, and beech forest, and also to the 
productivity of pastures. This seems, however, 
somewhat underestimated. From the growth 
analysis of a 12 year old stock of ash in Denmark, 
Boysen Jensen (17) estimated the annual produc- 
tion of dry matter per hectare as follows: 

1. Wood 4.09 tons 

2. Leaf 2.72 

3. Branches shed off 0.57 

4. Respiration loss 3.10 

5. Total 10.48 

Underground parts were excluded from the 
estimation. Assuming the Top/Root ratio to be 
5.0, annual net production (1+2 + 3) and gross 
production (5) may amount to 8.9 and 12.6 t/ha 
respectively. Miller's comprehensive work (18) 
clarified the age-productivity relationship in 
Danish beech forest. The net productivity reaches 


its maximum (ca. 16 t/ha/year) at 30-40 years, 
whereas the maximum gross production (ca. 28 
t/ha/year) is found at 60-90 years when increased 
ratio of respiration loss results in the decrease of 
net production by 2-4 t/ha. 

In 1956, Satoo (19) published the result of his 
growth analysis on a regenerated stand of aspen 
in central Hokkaido about 40 years in age, and 
estimated the net productivity of aerial organs to 
be 8.7 t/ha/year. With the assumption that 
ca. 30% of gross production is lost by respiration 
and T/R ratio is equal to 5.0, the estimated annual 
gross production becomes 14.9 t/ha. If the under- 
growth of bamboo grass is included in the cal- 
culation, it would rise to as much as 20 t/ha. 

Both Japanese and Danish estimates are on 
the same order of magnitude. Because Denmark 
and Hokkaido belong to the cool temperate zone 


No. of individuals per species 


Fig. "Z. Species number-individual number relation in a lucidophyllous forest of central Japan, dominated by Shiia 
cuspidata. The curve indicates Fisher's logarithmic series fitted to the data. Sampled area 0.16 ha. Data after Miki, 1932. 











"ioo iso 

No. of individuals per species 

Fig. 3. Species number-individual number relation in a tropical rain forest, Para, Brazil. Sampled area 3.5 ha. Data 
after Pires et ai, 1953. 




(warmth index approximately 50 and 65 month- 
degrees respectively), we may tentatively conclude 
that cool temperate forests produce 15-25 tons 
of dry organic matter per hectare and per year 
under moderate edaphic conditions. 

No exact estimates arc now available for 
warmer climates, but the production of some 
temperate grasslands may be worth mentioning. 
Although the standing crop of living plant of 
forest ecosystem greatly exceeds that of grassland, 
there is little evidence that the productivity of the 
latter is much lower than that of the former when 
they thrive under the same climate. In hilly 
pastures of Japan belonging to the temperate 
zone, where the natural grasslands, of Miscanihus 
sinensis are maintained for thatching materials, 

60 r 



a standing crop of summer foliage of more than 
60 tons per hectare (fresh weight) is common. 
The approximate equivalent of this in total dry 
matter including root is 27 tons. As the annual 
increment of living rhizomes seems negligibly 
small in a well-developed grassland, this amount 
of summer crop may roughly equal the net pro- 
duction of the year. Thus the gross productivity 
would amount to 39 t/ha/year. 

In this connection, the relation of agricultural 
productivity to thermal gradient is very suggestive. 
Kawakita (20) compared the agricultural pro- 
ductivity of eight districts of the Japanese Archi- 
pelago in terms of the average yields in calories 
of edible parts of main food crops. When 
the productivity is correlated with the average 

O O 







100 200 

Warmth index (month -degrees) 

Fig. 4. Fisher's index of diversity (a) increases with temperature. 






20xl0 3 









Mean annual temp. 

Warmth index 


200 month -deg. 

Fig. 5. The relation of agricultural productivity to thermal climate. 1: Sakhalin. 2: Hokkaido. 3: Northeastern 
Hondo. 4: Central and southwestern Hondo. 5: Sikoku. 6: Kyusyu. 7: Ryukyus. 8: Formosa. Data after Kawakita 


thermal data, approximately linear regression is 
obtained (Fig. 5). It is to be noted that the 
warmth index appears more linearly correlated 
to the productivity than the mean annual tem- 
perature. If similar linear regression be expected 
between the warmth index and the productivity 
of natural ecosystems and the gradient of the 
regression line remains as it is, we could estimate 
the productivity of tropical forests by extrapola- 
tion. According to the regression of Fig. 5, the 
productivity at 260 month-degrees of warmth 
index which is the standard value for humid 
equatorial climate is about 5.26 times as great as 
that at 55 month-degrees. Assuming the gross 
productivity of a forest at 55 month-degrees to be 
15-25 t/ha/year, the expected value in the tropics 
would be 79-132 t/ha/year, falling on the same 
order with that of Vageler's estimation. It is 
also of interest to note that the gross productivity 
of marine algae in a coral reef community of 
tropical Pacific water was 24g/cm 2 /day in glucose 
(21). Equivalent dry matter of this as ordinary 
plant materials is about 80 t/ha/year. 

A most reasonable representation of productiv- 
ity is found in the photosynthetic energy effi- 
ciency or the ratio of radient energy utilized for 
the organic matter synthesis to the total radient 
energy available. The efficiency or Lindeman's 
ratio has been studied mostly with aquatic 
ecosystems, and it is now widely recognized that 
eutrophic lakes and ocean waters in the temperate 
zone having phytoplankton as principle producer 
have the annual efficiency of 0.3-0.4%. Higher 
efficiencies were recently observed by Odum (21, 
22) in the tropical coral reef and the water-grass 
community of a freshwater thermostatic spring in 
Florida, the ratio to the total available radiation 
at the water surface being 2.9% and 2.6% res- 
pectively. One author (23) has stated that the 
energy efficiency in natural ecosystems hardly 
exceeds 1 % and that forests are in general even 
lower than aquatic ecosystems in their efficiency. 
However, our calculations on the aspen forest 
of Hokkaido mentioned above resulted in a con- 
siderably higher percentage. Annual gross pro- 
duction of 20 t/ha in this forest corresponds to 
the annual efficiency of 0.8%; and, if only the 
radiation during 5.5 months' growing period is 
considered, the ratio increases to 1.4%. As for 
the more productive beech forest of Denmark, 
where the available radiation becomes less owing 
to higher latitude (56N.), a still higher percentage 
is to be expected. For Miscanthus grassland of 
central Japan, the estimated efficiency is 1.9%. 
We might therefore conclude from these evidences 
that the forest ecosystem could reach a much 


higher level of energy efficiency than formerly 
believed, presumably as high as 2.5%. 

Average annual amount of solar radiation on 
the equator is 0.339 cal/cm 2 /min according to 
Simpson (24). This is equivalent to 1782 x 10 10 
cal/ha/year. Similar estimates by Fukui (25) 
give almost the same value. If the tropical forest 
is assumed to produce photosynthetic assimilates 
with the efficiency of 2.5%, 44.5 x 10 l calories 
are expected to be utilized for organic matter 
synthesis, or 117 tons of glucose are produced 
per hectare and per year. One hundred and seven- 
teen tons of glucose roughly correspond to 106 
tons of dry plant materials. This estimate again 
agrees well with the result obtained by the ex- 
trapolation of the productivity-temperature re- 
gression in Fig. 5. 


One of the greatest difficulties confronting 
tropical agriculture is the catastrophic decline of 
productivity that occurs when the forests have 
been cleared and converted into such artificial 
ecosystems as farmland or plantation. This is 
especially the case with humid rain forest areas. 
In general the agricultural exploitation of these 
areas is as yet far from success. Only the shifting 
agriculture on upland slopes and the paddy rice 
cultivation on irrigated plains, both of which 
have presumably originated among the inhabi- 
tants of humid tropics, are well adapted to this 

In temperate regions, the agricultural pro- 
duction is no less efficient than the production of 
natural ecosystems. Transeau (26) estimated, for 
example, the energy efficiency of gross organic 
matter production for a heavy crop of maize in 
Ohio to be 1.6%. Similar efficiency for average 
rice crop in Japan (ca. 4.6 tons of unhulled grains 
per hectare in air-dry weight) is about 1.1%. The 
highest yield of rice hitherto recorded from Japan 
amounts to 15.3 t/ha, and the gross productivity 
equivalent to this grain yield means the energy 
efficiency of about 3.3 %. Such high productivity 
of temperate agriculture is of course maintained 
by the addition of chemical and organic ferti- 
lizers, which not only compensates the amount 
of minerals removed from the soil as crops and 
through leaching, but also enriches the soil beyond 
the level of mineral content under natural condi- 
tions. Natural ecosystems, especially the forest 
with its stratified structure, seem more efficient 
in utilizing the available space as well as the 



radiant energy that falls upon it, as compared 
with singled-layered, widely spaced crop com- 
munities. It is perhaps the effect of enriched soil 
minerals that enables agricultural ecosystems to 
overcome this handicap. 

In the tropics, however, the return of minerals 
to the soil is so large in amount that to replace it 
with manuring under cultivated conditions is 
a hard task. On the other hand, high temperature 
accelerates the decomposition of litter to such an 
extent and the leaching of soil minerals is so rapid 
owing to heavy rainfall that the soil of cleared 
forest land is quickly devastated. We have seen 
in Fig. 5 that the agricultural productivity is 
nearly proportional to the warmth index through- 
out the long stretch of Japanese Archipelago, 
from Sakhalin to the Ryukyus, where the way of 
agricultural land utilization is essentially the same. 
Turning to Formosa, however, the productivity 
suddenly falls below the expected value. Al- 
though the change in the traditional system of 
land utilization may be responsible for the fact 
to a certain extent, this appears to suggest an 
unavoidable limitation imposed upon the tem- 
perate way of agriculture under tropical climates. 

Extremely low mineral content of tropical 
forest soils must be mentioned here. In regions 
of nearly equal climatic humidity, the organic 
matter or the nitrogen content in the soil can be 
represented as the function of temperature, 
decreasing exponentially with rising temperature 
(27). In contrast to temperate forest soils con- 
taining an average of 0.1-0.3% total nitrogen, 
the percentage for tropical forest soils hardly 
exceeds 0.05%. Similar relations may well be 
expected for other mineral nutrients. The layer 
of undercomposed organic materials is also 
poorly developed on the floor of tropical rain 
forest. But the minor standing crop of minerals 
stored in the soil does not always indicate the 
lower productivity of a whole ecosystem, as the 
standing crop is nothing but the balance in the 
budget of minerals which at every moment are 
set free from the litter and absorbed by the roots 
again. Several authors have reached the same 
conclusion (14) that there is an almost closed 
cycle of plant nutrients in the tropical rain forest 
ecosystem and that the minerals are at once 
reabsorbed by plants as soon as they are released 
from the decomposing plant materials. 

The dynamics of soil mineral budget is quite 
different in temperate forests. To cite an example 
from the study of Ovington (28), average dis- 
tribution of nitrogen in several pure stands of 
different tree species, artificially established at 


Abbotswood, England, were as follows: 

In canopy 274 kg/ha 

Tree boles 132 

Ground flora 32 

Forest floor 235 

Mineral soil 7,041 

(Underground parts are omitted) 

Considering the age of stands (mostly 40-45 
years), the amount of nitrogen in the annual 
increment of living plant body as canopy, boles, 
and ground vegetation may be of the order of 
200-250 kg/ha. When compared with the amount 
present in soil and litter (235 + 7,041 - 7,276 
kg/ha), the turnover of soil nitrogen would be 
only 1/30-1/35 time/year, if the addition due to 
rainfall and nitrogen-fixers be compensated by 
the loss through leaching. 

As to tropical forests, judging from the pro- 
ductivity and the average nitrogen content of soil, 
the annual turnover of soil nitrogen might be not 
far from 1. Namely, the speed of circulation of 
plant nutrients is 30-35 times more rapid in trop- 
ical than in temperate forest. The speed of 
circulation is most probably the function of 
temperature, and to establish the turnover-tem- 
perature relation as known functions may provide 
a possible approach to understand the dynamics 
of the tropical rain forest ecosystem. 

What we have said above is little more than 
common ecological knowledge. Our intention 
is to point out the importance of the study of 
ecosystem metabolism in the rain forest and its 
implications to tropical agriculture. Intensive 
study in forest areas where shifting cultivation is 
prevalent may throw new light on the problem. 


1. It is pointed out that the tropical rain forest 
occupies a terminal situation on a long continuous 
series of forest vegetation from subarctic to tro- 
pical climate. Along the scries or the latitudinal 
thermal gradient on the earth, the floristic diversi- 
ty, productivity, and other characters of forest 
ecosystem continuously changes, ending in the 
tropical rain forest with its extreme richness in 
flora and organic matter production. The im- 
portance of the western Pacific area is stressed 
which represents the sole example in the world of 
uninterrupted continuation of humid forest 
climate from high latitudes to the equator. 

2. By the extrapolation of a temperature- 
productivity curve obtained from temperate 
regions, the annual gross productivity of tropical 

rain forest is estimated to be 105 25 t/ha in dry 
matter. This agrees well with another estimate 
based on the energy efficiency of organic matter 
production by forest combined with the amount 
of total radiant energy available in the tropics. 

3. The cause of the failure of rain forest area 
as arable land is discussed as related to the eco- 
system metabolism. It is suggested that the very 
rapid circulation of plant nutrients between plant 
and soil is mainly responsible for the sudden 
decline of productivity that ensue from the clear- 
ance of rain forest. A rough estimation has 
shown that the turnover of soil minerals may be 
some 30 times greater in tropical forest than in 
temperate forest. 

4. These results are only tentative, but they 
may serve as the starting point for intensive field 
researches on tropical rain forest which are 
earnestly desired in the near future. 


(/)* Kira, T., 1945, A new system of climatic 
classification in eastern Asia as the basis 
for agricultural geography, Kyoto. 

(2)* Kira, T., 1945, New classification of cli- 
mates in southeastern Asia and western 
Pacific, Kyoto. 

(3)* Suzuki, T., 1952, The East-Asiatic forest 
climaxes, Tokyo. 

(4) * Imanishi, K., 1953, Nature of Nepal Hima- 

laya. Kagaku 23:406-416, 464-468. 

(5) Kawakita, J., 1956, Vegetation. In 'Land 

and crops of Nepal Himalaya', Scientific 
results of the Japanese expeditions to 
Nepal Himalaya, 1952-1953, edited by 
H. Kihara, Tokyo, pp. 1-65. 

(6)* Imanishi, K., 1937, Community analysis 
and community classification. Geogr. 
Rev. Jap., 13:725-736. 

(7)* Imanishi, K., 1937, The altitudinal regions 
of the Northern Japanese, Alps. Jour. 
Jap. A/pine Club, 31:269-364. 

(8)* Imanishi, K., 1949, Principles of bio- 
sociology, Osaka. 

(9) Curtis, J.T. and Mclntosh R.P., 1950, An 
upland forest continuum in the prairie- 
forest border region of Wisconsin, 
Ecology, 32:476-496. 

(10) Brown, R.T. and Curtis, J.T., 1952, The 
upland conifer-hardwood forests of 

* In Japanese. 

** In Japanese with English summary. 


northern Wisconsin, EcoL Monogr., 

(11) Whittaker, R.H., 1952, A study of summer 

foliage insect communities in the Great 
Smoky Mountains, Ecol. Monogr., 22: 

(12) Whittaker, R.H., 1956, Vegetation of the 

Great Smoky Mountains. EcoL Monogr., 
26: 1-80. 


Fisher, R.A., Corbet, A.S. and Williams, 
C.B., 1943, The relation between the 
number of species and the number of 
individuals in a random sample of an 
animal population, Jour. Animal 





Richards, P.W., 1952, The tropical rain 

forest, Cambridge. 
Vegeler, P., 1930, Grundrissdertropischen 

und subtropischen Bodenkunde, Berlin. 
Filzer, P., 1951, Die naturlichen Grund- 

lagen des Pflanzenertrages in Mittel- 

curopa, Stuttgart. 
Boyscn Jensen, P., 1932, Die Stoflproduk- 

tion der Pflanzen, Jena. 
Boysen Jensen, P., 1949, Causal plant geo- 
graphy, Del Kgl. Danske Videnskah. 

Selskab, Biol. MeJd., 21 (3): 1-19. 

(79j**Satoo, T., Kunugi R. and Kumekawa, A., 
1956, Amount of leaves and production 
of wood in an aspen (Popitttts Davidiana) 
second growth in Hokkaido, Bull. Tokyo 
Univ. Forests, 52: 33-51. 

(20)* Kawakita, J., 1949, A quantitative repre- 
sentation of land productivity in terms 
of calory yield. Geography for Social 
Life, 19: 6-10. 

(21) Odum, H.T. and Odum, E.P., 1955, Troph- 

ic structure and productivity of a 
windward coral reef community on 
Eniwetok Atoll, EcoL Monogr., 25: 

(22) Odum, H.T., 1957, Trophic structure and 

productivity of Silver Springs, Florida. 
EcoL Monogr., 27:55-112. 

(23) Park, O., 1949, Community metabolism. 

In Allee, W.C. ct al. : Principles of 
animal ecology, New York. pp. 495-528. 

(24)* Fukui, E., 1941, Climatology. Revised 

Edition, Tokyo. 
(25)** Fukui, E., 1953, Climatology of radiation; 



the meridional distribution of insolation (27) Jenny, H., 1941, Factors of soil formation, 

over the earth's surface. II., Geogr. Rev. New York. 

Jap., 26: 573-585. ^ Ovington, J.D., 1957, The volatile matter, 

(26) Transeau, E.N., 1926, The accumulation organic carbon and nitrogen contents 

of energy by plants, Ohio Jour. Sci., of tree species grown in close stands. 

26: 1-10. New Phytologist, 56: 1-11. 






Botanical Institute, Hiroshima University, Hiroshima, Japan. 


As one of the most characteristic features of 
the Japanese flora, it is cited that several plants of 
the tropical origin are growing wild there. This 
fact was already clarified by me with respect to 
bryophytes. As to the vascular plants, the follow- 
ing is known about the distribution of some 
species. According to their northernmost limit 
of habitat in the Japanese Archipelago, the species 
are divided into ten types: 

Type 1 . ranging from Indo-Malaysia north 

to 44 L.N. in Japan. 

Trichomanes parvulum Poir. 
Type 2. to 41 L.N. 

Vitex rotundifolia Linn, fil., Albiz- 

zia julibrissin Durazz. 
Type 3. .__to 38 L.N. 

Dicranopteris glauca Underw., Hy- 

menophyllum bar bat um Baker, Dro- 

sera spathulata LaBill. 

Type 4. to 37 L.N. 

Psiloswn nudum Beauv. 

Type 5. _to 36 L.N. 

Ceratopteris thalictroides Brong., 
Mecodium poly ant has Copel., Ly- 
copodium cernuum Linn., Vittaria 
flexuosa F6e. Anodendron affine 
(Hook, et Arn.) Druce. 

to 35 L.N. 

Nephrolepis cordifolia Presl., Ipo- 
moea indica (Burm.) Merr. 

Type 6. - 

Type 7. __ 


Histiopteris incisa J. Smith, Humata 
repens Diels, Cocculus laurifolius 
DC, Solarium aculeatissimum Jacq., 

Type 8. 

Dichondra repens Forst, Ipomoea 
pes-caprae (Linn.) Sweet, Senecio 
scandens Hamilt. ex D. Don, 
Dianella ensifolia (Linn.) DC. 33 L.N. 
rnacrorrhiza (Linn.) 


Type 9. . ... . to 31 L.N. 

Ficus retusa Linn., Kandelia Candel 
(Linn) Merr., Messerschmidia ar- 
gentea (Linn., fil.) Johnston. 

Type 10. to 30 L.N. 

Blechnopsis oriental is Presl., Bias- 
tus cochinchinensis Lour., Melas- 
toma candidum D. Don., Spinifex 
Uttoreus (Burm. fil.) Merr. 

(The horizontal and vertical ranges of each 
species are to be discussed with the frequency 

At the present stage of our knowledge, it is 
hardly possible to form a safe conclusion about 
the phytogeographical explanation of the occur- 
rence of these tropical plants in the Japanese Ar- 
chipelago. It is more likely that these species 
were once distributed much more widely in tem- 
perate regions of East Asia, such as China, Korea, 
and Japan. Jn later times, through climatic 
change, they were unable to compete with more 
aggressive species and have been reduced to their 
minor role in the vegetation. The coast of the 
islands of Japan has been warmer under the 
influence of the warm sea-current of "Kuro- 
shio," and they may have survived here as living 





A ton Forest, Norfolk, Connecticut, U.S.A. 

My role in summarizing this symposium on 
Vegetation Types of the Pacific Basin is fraught 
both with customary and uncustomary difficulties. 
J should state first that my position as Chairman 
of this program is only as a latter-day substitute 
for Dr. Pierre Dansereau, to whom should go full 
credit for organization, and who regrettably was 
unable to come to Bangkok despite earlier plans 
to the contrary. 

During the course of this symposium we have 
been presented with several succinct and admira- 
ble contributions. At first glance, it may appear 
that these are totally disconnected and unrelated 
"islands" in the "sea" of vegetation (plant com- 
munity) science. Perhaps we should face one fact 
at once: the terrestrial vegetation of the Pacific 
Basin, as distinct from that of the rest of the 
world, has but one element that binds it all 
together a vast mass of water called the Pacific 
Ocean which is not a substratum for terrestrial 
vegetation, and acts as the best possible agent to 
prevent the unification of that vegetation. Thus 
the Pacific Basin is a geographic entity, united 
more by the common interests of its human popu- 
lations than by any fundamental vegetational 

The papers presented in this symposium serve to 
place the spotlight on a few elements of the whole 
problem, much as if a spotlight were to pass over 
a crowded stage, temporarily lighting a few per- 
sonalities, but leaving all the rest in darkness. 
This momentary lighting however is very im- 
portant. It serves to add a definite increment to 
our knowledge of Pacific Basin vegetation, even 
as does each succeeding Pacific Science Congress. 
My purpose at this time is no more than to place 
these papers in the larger framework of such a 

There are several "viewpoints" from which 
vegetation may be approached. No one view- 
point has precedence over any other. In the 
rest of botany, for example, the morphologist 
does not vie with the physiologist as to which is 
most "important," or which comes "first." So it 
should be with the students of plant communi- 
ties. The floristic composition of vegetation is 
one such viewpoint. Within the Pacific Basin, 


the floras segregate into at least three majoi 
groups. There is a circum-Arctic flora on the 
mountains near the Alaska coast, out across the 
Aleutians, and down the high elevations of the 
Kamchatka Peninsula. This flora, giving rise tc 
tundra types of vegetation, has not been repre- 
sented on this symposium. Progressing south- 
wards, we find temperate floras that have more in 
common with themselves than with the arctic 01 
tropical floras that bound them on each side. 
Becking, Tatewaki, and Chi-Wu Wang have each 
contributed to our knowledge of these areas, 
The tropics, by comparison, is a relatively unified 
region, characterized in its vegetation by the 
relative absence of damaging cold weather (al- 
though how that damage is described in purel} 
meteorologic terms is still an ecologic puzzler), 
Porleres, Hiirlimann, Kira, Horikawa, and Fos- 
berg have given us interesting information or 
vegetations developed from these floras. Of these. 
Fosberg has handled the atoll problem, where the 
flora is attenuated to point of poverty, thus 
greatly simplifying some aspects of vegetation 
study. The temperate floras of the southern he- 
misphere, of Australia, New Zealand, and South 
America are not represented on this symposium, 
except for Fosberg's comments on the montane 
grasslands of the Andes. 

Passing on to a second viewpoint, interest in 
the form and structure of vegetation is a major 
preoccupation of some vegetation scientists, 
Becking, in handling northwestern United States, 
is concerned largely with the make-up of vegeta- 
tion in terms of the plant communities, according 
to the ideas of Braun-Blanquet. Hurlimann is 
concerned with structural details of a tropical 
forest. Thirdly, Dansereau has presented us 
with an expression of his system of shorthand 
symbols for recording the structure of communi- 
ties. Like most systems of shorthand, it is be- 
wildering at first look, particularly this lollipop- 
oriented one. (Freudian overtones specificall> 
not implied!) Furthermore, Dansereau is ver> 
much alive, and I fear there may be revisions tc 
the revisions before the gentleman passes on tc 
a stable end-stage of heavenly climax, revisions 
that make it difficult for ones of limited mentalit) 

like myself to keep up with him. It was ques- 
tioned from the audience as to whether this 
system had been or could be used in the floristical- 
ly more complex tropics. It is my personal 
opinion that it is in areas of just such complexity 
where this system might prove most serviceable, 
in that it describes structure unrelated to details 
of floristic composition, which details for most 
tropical regions are not adequately known. 

One micro-facet of the morphology of plant 
communities is that of epiphyte communities, of 
those aggregations of plants which utilize another 
plant as a substratum, and which can be studied 
as a separate element in the total phytosociologic 
picture. Hosokawa and his colleagues have car- 
ried on a great many studies with epiphyte com- 
munities, both in Japan and on Pacific islands. 
Their present contribution is one more in a 
notable series. 

Another aspect of the form and structure of 
vegetation, related not to the different communi- 
ties involved, or to the structure of those commu- 
nities, but to whole mosaics of communities, is 
the grassland-savanna-forest problem. These 
types of vegetation are found in close association 
with each other in many parts of the tropics. 
Their interpretation as being "natural," or 
induced by primitive tribes and persisting into the 
present as "relicts," or induced by contemporary 
agricultural and pyric factors, is a phase of scienti- 
fic investigation discussed by Porteres and Fos- 
berg. It is a fascinating field of inquiry not only 
for the academic mind, but for the Vegetation 
Manager who wants to manipulate and convert 
vegetation, sometimes into types markedly differ- 
ent from what now exists. 

In the third "viewpoint," that of functions and 
processes, it is quite interesting that not a single 
paper is developed along the traditional lines of 
"plant succession" to "climax," with an orderly 
diagram showing the origins of the various 
"seres," culminating by means of arrows, in one 
final end-stage. Nevertheless, all the speakers 
have considered adequately and worthily the 
various dynamic functions that are involved in the 
life-activities of their various vegetations. 

The "ecologic" (environmental) approach, a 
fourth viewpoint, is coincidentally absent as 
the major approach in any one contribution to 
this symposium. This approach, dominant in the 
thinking of possibly the major part of vegetation 
research, at least in America, is not omitted from 
this symposium by intent. Although we have no 
paper entitled "The influence of climate on the 
distribution of . . ." or "Soil chemistry as related 


to the vegetation pattern in . . .," there has been 
no slight, with any speaker, as to an awareness 
of the importance of the environment in the 
behavior and phenomena of vegetation. 

The geographical distribution of vegetation 
types is a fifth viewpoint, and cartography (map- 
ping) is one of its most important aspects. In this 
branch of knowledge, Kuchler has distinguished 
himself, particularly in accumulating a world 
bibliography of vegetation maps. His presenta- 
tion and views serve excellently to round out 
this symposium. Aerial photography has enor- 
mously enhanced our facilities for vegetation 
mapping in recent years. It should be pointed out, 
however, that the vegetation types which are dis- 
tinguishable from the air are generally the phy- 
siognomic types, that is, types distinguished as 
forest, grassland, savanna, and desert. Equally 
distinguishable are types dominated by an aer- 
ially visible species, such as pine forests in a 
mixed hardwood region. There are dangers as 
well as advantages to this facility, for it leads us 
to overlook the fact that there may be other and 
even more significant kinds of vegetation types 
which are not related to these aerially visible 
differences. Therefore one cannot overemplasize 
the need for accessory studies on the ground by 
competent botanists. 

A sixth and final viewpoint, that of distribution 
in time, or history, has also been underplayed, 
if we view its importance in an ideal vegetation 
science. Both Porteres and Fosberg have 
touched the subject in their inquiry on the 
possibly-human origins of grasslands and savan- 
nas, and Chi-Wu Wang has considered the history 
of the Chinese forests as it extends back into 
geologic time. 

As a postlude in one sense, and as an open door 
to larger vistas in another sense, the spotlight 
should be played again on Fosberg's treatment 
of atoll vegetation. The idea implicit here is 
developed more extensively by him in another 
paper delivered elsewhere at this Congress, "... 
Description of the Coral Atoll Ecosystem." 
Now in all other papers of this Symposium, we 
have considered "vegetation" as our subject of 
study. For many years, however, scientists have 
realized that vegetation is but part of a larger 
integrated whole, called by such names as eco- 
system, microcosm, landscape, and "organism" 
(not the biologic organism). The obvious intellec- 
tual advantages of this approach are often offset 
by the very complexity of the natural phenomena 
involved, and the fact that adequate scientific 
knowledge from many disciplines is seldom 



encountered in one man. The atoll, by its very 
nature, is a distinctive microcosm, existing at the 
interface of sea and air, with energy absorbed 
from and radiated to its environment, with a 
sort of feed-back mechanism that keeps the 
system in a relatively steady state. From this 
view, the vegetation all but loses its separate 
identity in the structure and behavior of the larger 
"whole." It is not to be implied that the same 
conceptual approach is immediately applicable to 
continental areas, with their enormously greater 
complexities, and less clearly defined boundaries 
in space. Fosberg's researches greatly advance 
our ideas in these conceptual realms, and indicate 
that the views are both feasible and practicable. 

In closing this symposium, which had been 
planned by Dansereau as Chairman of the 

subcommittee concerned with Vegetation (plant 
communities), I wish to say that in my opinion 
this subcommittee, acting within the framework 
of these Congresses of the Pacific Science Associa- 
tion, has a brilliant future ahead of it. A unified 
vegetation science is just beginning to emerge 
from the various nationalistic disciplines which 
originated mainly in North America and Europe. 
There is now no other area in the world than this 
Pacific Basin where scientists of so many nations 
can cooperate so advantageously, with the vigor 
that comes from the interplay and interchange 
of ideas, of problems, and of research accom- 
plished. I look forward to seeing this vigor 
translated into a distinct world contribution 
towards the description and understanding of 
the world's vegetation. 




Flora Maletiana Foundation, Oegstgeest, Netherlands. 

A proper understanding of plant structure is 
an absolute necessity for botanical education, 
without which pupils of the secondary schools 
the source from which university students will be 
recruited will not be able to understand plant 
function, plant ecology, flower biology, forestry, 
and agriculture. 

The identification of a living plant by means of 
keys and a description is still the best means to 
gain an insight in its structure and its contrasting 
features with other plants. Plant atlasses may 
seem to lead to the same result along a more easy 
way. But this is educationally basically wrong, 
it leads only to a vague notion of the general 
habit of a plant and to its name, not to any knowl- 
edge of its structure. 

Most Floras have a dualistic character, they 
tend to give a complete survey of all the species 
of a country or a more restricted area and must 
serve simultaneously for education. 

In tropical countries this dualism is untenable 
by the wealth of the floras, and would necessitate 
large and expensive works in which the educative 
purpose is entirely lost. Such Floras would con- 
tain descriptions of plant forms from remote 
mountains and forests which could never serve 
for regular class demonstration and which no 
teacher could be expected to know himself, let 
alone have available in the living state in sufficient 
quantity for class teaching. 

Complete Floras may be desirable for botanists 
and advanced amateurs, but for school-children 
they represent the 'Tables of the Law', they have 
no educative value and are deterrent rather than 
awakening interest in plants. 

It is compulsory that in the class each pupil 
has in hand a living specimen and the Flora, with 
at most a pin or needle as a simple utensil; lens 
and forceps will generally be not available and 
are unnecessary for the purpose. 

The idea is that the identification of a plant is 
performed in the class, step by step, and that 
under guidance of the teacher each pupil can 
follow the way along which the ultimate goal, 
the name, is reached with subsequent checking 
of the description. This simple botanical detec- 
tive work must give satisfaction. 

Such a School-flora must be adapted to the 
following points: 

(a) A selection of plants, hence no complete- 
ness whatsoever. 

(b) Availability of at least a number of species 
incorporated in it, as the teacher must be able to 
collect for one lesson enough material to give a 
specimen to each pupil. Consequently the choice 
of selection is restricted to plants available in or 
near towns and cities. 

(c) Extensive use should be made of cultivated 
plants from school-gardens or nurseries, town 
parks, roadsides, etc., resulting in inserting in the 
Flora numerous cultivated and ornamental plants. 

(d) To bring pupils into contact with biotopes 
in his immediate vicinity brings along the insert- 
ing in the Flora of some typical plants of beach, 
mangrove, rice-field, swamps or pools, dry- 
farming fields, estates, waste ground, secondary 
growths, etc. 

(e) The absence of a lens and forceps and 
knowledge how to use these necessitates the 
omission of plants with small flowers which 
escape easy observation with the naked eye. 
Besides, there is no educative advantage in dis- 
secting small flowers against using large ones. 

(f) There must be no obscurity in the identifi- 
cation keys to the family, genus or species, and 
when the name of the latter is reached there 
should be a clear not too elaborate description for 
checking the identity. Great care should be given 
to attain a faultless text. 

(g) Interest in the plant should be raised in 
adding where desirable some brief additional 
notes on uses for mankind, biological topics, or 
other noteworthy data. 

(h) In absence of adequate pre-knowledge 
technical terms should be avoided and the keys 
and descriptions should be as much as possible 
worded in plain language ; a plate showing schema- 
tic pictures to elucidate necessary technical terms 
may be handy. 

(i) Representatives of the most important 
tropical families should be inserted by native, 
introduced or cultivated species. 



With such a flora only a select number of species 
can be identified; it serves merely for educative 
purposes', the idea of completeness is entirely 

If we imagine that during the courses in the 
secondary school not more than about 25-50 
species will be identified in the class, the School- 
flora should contain not more than about 300- 
400 species in all. 

With a few collaborators I have realized such 
an octavo School-flora for Indonesia, with about 
400 species belonging to about 300 genera and 
120 families. It covers about 400 printed pages 1 . 

It is not the intention that pupils will know or 
learn all these 400 plant species; no more than 
10% can be treated in the class. But the number 
of 400 leaves room to the teacher to make his 
choice in accordance with available plants in 
sufficient quantity in his vicinity. Representative 
species of both the everwet and seasonal regions 
should be inserted. It further allows pupils who 
are interested to use the book for their own 

School-floras, like Alston's The Kandy Flora' 
(1938) and Merrill's 'Flora of Manila' (1912) 
follow a scheme which is less adapted to practical 
use as our School-flora for Indonesia. Both 
contain generic diagnoses which I feel are un- 
necessary for secondary schools. 

My main objection is, however, that they re- 
present a complete flora of the vicinity of a town, 
Kandy and Manila respectively, without restric- 
tion to easily available plants for class teaching 
and selection on size of flowers. They both still 
contain a dualistic element (serving education and 
botany) and are not entirely devoted to the goal 
of being merely a tool for educative purposes. 

In the School-flora for Indonesia we could not 
avoid mention of such important groups as 
grasses, sedges, and ferns, and have been com- 
pelled to add some simple pictures for a few re- 
presentatives of these groups. 

The insertion of common and widely distributed 
ornamentals, weeds, fruit trees, wayside plants, 
etc. is convenient to use the flora in the whole of 
Indonesia in cities and towns below 1,000 m 
altitude, the site of most training centres. And 
I am rather certain that it could for the same 
reason be used in most parts of tropical southeast 
Asia, Micronesia, and Melanesia. 

I believe that the model of a School-flora of 
which the principles have been just outlined, is 
a valuable tool for education purposes and the 
wakening of the interest of pupils. 

I find it desirable to bring these principles to 
the general notice of teachers and hope it will be 
a stimulant to the writing of similarly adapted 
School-floras in other tropical countries. 


F.R. FOSBERG: E.H. Bryan, of the Bishop Museum is 
writing the flora of Guam which fills Van Stccnis require- 
ment for School-flora. The common species are empha- 
sized so that the ordinary people of Guam can learn. Such 
books acquaint young students with the nucleus of botany 
before college time. 

C.G.GJ. VAN STEENIS: In Java, education in schools is 
faced with the problem that knowledge of Java's flora 
(4,500 species) is impossible. Therefore it is a necessity 
to have easy tools to instruct people in the morphology of 
certain plants. One disadvantage is that the teacher must 
know plants and names himself. 

M.L. STEINER i I do not agree with the exclusion of small 
species of flowers. 

C.G.G.J. VAN STEENIS: For secondary classes it is suffi- 
cient to study only large flowers. 

M.H. SACHET: I disagree. 

F.R. FOSBERG: People may learn botany, but not of 
plants. America has a tendency not toward School-flora 
but toward picture books which form bad habits. 

p. WEATHER WAX: A written examination would find 
out if students can use the key. 

V.A. JIRAWONGSE: Is there an advantage in using just 
vernacular names? 

C.G.GJ. VAN STEFNIS: Vernacular names are different 
in different parts of the world, and Latin names are used. 

"Flora voor de scholen in Indonesia". Noordhoff-Kolff, Djakarta, 1st ed. 1949, 2nd ed. 1951. 






Faculty of Agriculture, Pajakumbuh, West Sumatra, 


Sumatra, as compared with Java, is a land of 
promise for botanical exploration. On the 
densely populated island of Java, most of the 
lowland flora is destroyed, and almost the whole 
country is divided among agriculture, forestry, 
cattle-breeding, and fishery. Botanical explora- 
tion has already been so thorough in Java that 
only rarely are new plants found. Bogor is still an 
important botanical centre on Java. In Sumatra 
we find a quite different situation though not 
everywhere. It is not only a land of promise but 
also of contrasts. Regions like Djambi and 
Indragiri are still largely covered by primary 
forests in which are found elephants, rhinos, and 
tigers. Only 2-5 plant specimens have been 
collected per 100 sq km in these regions. I read 
in the newspaper that the tribes of the primitive 
Kubu-people are still living in these forests. It is 
told that they walk undressed and are forbidden 
by their own law to plant. 

Quite different situations are found in other 
places: the Deli-region in the Northeast has been 
widely cultivated and is still the area of big estates 
of oil palm and tobacco and rubber. Minangka- 
bau on the West Coast is to be seen from the air 
like an enclave between the jungles of South 
Tapanuli, Indragiri, Djambi, and Bengkulu. It 
was long ago colonized by people who made this 
a region one of mixed garden agriculture and who 
still preserve their special customs. The land- 
scape has much the same features as Java, but 
agriculture is far less intensive than in Java 
and the flora of the kampongs also shows many 
differences. Minangkabau people possess, for 
example, their own races of rice (hundreds!); 
their own names for and assortment of bamboos ; 
their own names for at least twenty-five different 
kinds of cultivated bananas; their own way of 
planting tobacco, cinnamon, sugarcane, clove 
(tjenkeh), and gambir; their own way of burning 
grassfields and secondary forests, of plundering 
the forests, taking no care of silviculture, land 
and soil protection. It is very difficult to change 
cultivation customs which cause destruction of 

t Presented by J.V. Santos. 

the landscape and impede further development. 
Sumatran students entering the new University 
Andalas are not selected among the Kubus or 
among kampongs far from the town; they are 
largely the sons and daughters of businessmen 
and government officials urbanized people who 
have lost contact with their rural surroundings. 
They have been strongly influenced by the 
blessings of modern culture from America 
and Europe, but are still in search of their own 

Pajakumbuh, is an ideal spot for botanical 
exploration and education. Here we find in a 
region of 10 sq miles (about 14 km square), 
a flat basin with mixed gardening and marshes 
(Pajakumbuh means the plain of marshes), 
surrounded by hills on which tobacco and gam- 
bir are planted and forest clad mountains com- 
posed of three different types of rocks: sandstone, 
limestone, and the old volcano Mt. Sago. The 
lowland forests, rich in Dipterocarpaceae, of the 
large East Sumatra coastal region are on a dis- 
tance of about 1 5 miles. The whole flora of the 
Pajakumbuh region probably has at least 2,000 
species of flowering plants. For Mt. Sago 
alone, I could list more than 800 species. The 
whole phanerogamic flora of Sumatra may be 
estimated as about 10,000 species including, for 
example, 800 orchids. 

When we read in Flora Malcsiana that the 
collecting density in W. Sumatra per 100 sq km 
is 38 (332 in W. Java), we can understand why 
our recent collections from the Pajakumbuh 
region contain numerous novelties new to 
science, or plants hitherto known only in Malaya, 
or collected only in South or North Sumatra. 
Several of these plants are really common in 
special types of vegetation. It is becoming clear 
now that forest types are strongly correlated with 
soil types, the so-called climax theory has no 
scientific basis! The forest vegetation on lime- 
stone hills is quite different from that on sand- 
stone hills. Analogous differences are seen in the 
weed vegetation and the cultivated races of rice. 
Limestone regions of the plain are very different 
from sandstone regions. The latter appeared to 
me in need of phosphate manure which could be 



found in caves in the limestone hills! We are 
drawing the attention of the agriculture service 
to this situation. 

We have already more or less compared the 
flora of Mt. Sago with several other mountains 
in Central Sumatra. Impressions of the lowland 
forests were collected in West-Indragiri. The 
sandstone region was the subject of the most 
recent expedition. More thorough studies of 
the limestone flora and of marsh forests remain 
for the future. We are presently making a 
thorough inventory of economic plants in the 
Pajakumbuh region devoting attention to all kinds 
of fruit trees, vegetables, legumes, races of rice, 
weeds, and secondary vegetation. We are able 
to advise and help the population in the rural 
reconstruction area served by our Faculty to 
use their waste lands more efficiently and to 
improve their mixed gardens. The culture region 
is far less productive as compared with Java. 
Our hope is that investigation will precede 
long-run planning in Central Sumatra and that 
natural resources will be better preserved than at 
present. Reforestation should become more 
than incidental planting of Pinus, Acacia auri- 
culaeformis, and some Toona sinensis. Good 
measures against burning and for forest and soil- 
protection should be taken. The farmers ought 
to be educated in this direction. 

We are ourselves trying to do this in our recon- 
struction area. We have succeeded in getting 
the road improved and have gathered all kinds of 
informations about economic botany, started a 
reforestation project and have built a nursery- 
garden. We are now engaged in project to plant 
the waste hills which are burned every year and 
show heavy signs of erosion. Our exploration task 
cannot be separated from our informations and 
education. The local population needs much 
information and education. People are no 
longer content with their standard of life. Indone- 
sian society is in transition! Descriptive botany 
and botanical ecology are very useful and practi- 
cal in tropical agriculture ! 


There will be no practical application of our 
studies in forestry, cattle-breeding, fishery and 
agriculture because they have no role in the 
botanical education of the country. Technical 
assistance for this requires more than three years. 
The Dean of our Faculty estimates that ten years 
will be necessary to get Sumatra's own people 

sufficiently interested and educated in scientific 

Are the facilities for such an education already 
provided at the new University of Andalas ? No ; 
after two years of struggle with many kinds of 
difficulties, a botanical institute is being built, but 
because of a lack of equipment, it is still impossible 
for the departments of plantphysiology, micro- 
biology, and cytogenetics to do their work on 
a modern level. Our Faculty is still without any 
laboratories for physics, chemistry, and agro- 
nomy. For social-economic science we have 
nobody! In Indonesia it often takes two years 
before ordered instruments arrive. The education 
of skilled personnel takes much patience and 
good-will. Minangkabau people have another 
discipline of work than Europeans or Americans. 
Foreign medical doctors feel the same difficulties, 
but fortunately self-education and self-criticism 
are also known here. The non-experimental 
parts of botany can start from the first days of 
arrival of a field botanist. Old newspapers for 
drying plants are to be bought on the local 
market; bamboo sasaks are easily made; and the 
sun is the great drying oven: Kampong people 
may be found who know the roads into the forests 
and many local tree names. The botanist must 
be able to get the authorities interested in his work ; 
he needs transport, funds, and even soil for expe- 
rimental gardens. Knowledge of the Indonesian 
language to educate the students in botany is 
very helpful. A good exploration of the nearby 
kampongs and mountains makes it possible to 
teach botany outside the lecture room in its 
natural surrounding. Then you have the simple 
task of showing the new students for the first time 
in their life what cotyledons really are (the word 
Dicotyledons they know) ; opening their eyes to the 
special structure of flowers of coconut palm, 
banana, and papaya; teaching them to distinguish 
fruit trees, legumes, vegetables, curbits, green- 
manures, and ornamentel plants; teaching them 
to observe plant forms, to describe them, and to 
learn how to make use of keys for identification. 
It is astonishing how little is known about this; 
a lot of goodwill, patience and perseverance is 
asked from the teacher. As soon as microscopes 
Arrive in my case after one year and the funda- 
mentals of plantmorphology and knowledge of 
plants in the surroundings are understood, you 
may start with laboratory courses in plant anato- 
my and cryptogams. Please do not look for Rhoea 
discolor to demonstrate plasmolysis; use the red 
leaved Cannas one meter from your door. As to 
anatomy, don't look only for the classical exam- 
ples, but take Portulacca, Sida, and Amaranthus 



growing along the door. The classical systems 
of vascular bundles without secondary growth 
seem to be an exception among tropical plants. 
Show the students who know only red and white 
rice, the hundreds of varieties growing in the 
surroundings. They will be astonished, enjoying 
the wide range of variation like every taxonomist 
enjoys seeing how nature plays with forms. 

Make the students aware of the fact that their 
own country should be the real centre for cultiva- 
tion and breeding of bananas, not Central or 
South America. Eradicate the theory that bana- 
nas were imported from America into Indonesia; 
eradicate the idea that the clove came from 
Zanzibar, that the local Rafflesia the biggest 
flower of the world is insectivorous. Demon- 
strate the wild species of Musa growing on 
Mt. Sago how you describe them and document 
your finds with photographs, how you further 
investigate the many cultivated varieties. Don't 
teach them a mass of facts, but methods and 
synthetic views. What a difference between an 
ordinary paddy-field seen as a basin with rice or 
seen as a dynamic whole, a habitat in and 
around which phanerogams, weeds, algae, bac- 
teria, soil composition, and the origin and 
breeding of rice-varieties all play a role ! What a 
difference between a kampong as an ordinary 
collection of fruit trees and vegetables and the 
same seen through the vision of a botanist who 
points out the origin of some elements in the 
forests which grew here during former times; 
the influence from India, from America; the 
influence of Dutch agriculture; the richness in 
varieties, the flower and seed biology and 
embryology of Manggas, Djeruk, Djambu, and 
Bananas; the role played by algae and water- 
plants in small fish ponds fertilised from the 
primitive houselets above them. 

The most clever and interested students may 
take a part in expeditions to the highest mountain 
summit of Sumatra (Kerinchi, 3,800 m) or in the 
lowland forest of Indragiri. They learn the most 
common forest plants; they learn how to organ- 
ize an expedition, how to collect plants, how to 
documentate the collection. They become 
teachers in secondary schools, discuss with you 
during evenings in the bivaques the best system 
for teaching botany there and arrange for you to 
give popular lectures to the local population, 
pupils of secondary schools, and their teachers. 
It might be necessary then to borrow clean trou- 
sers from one of your students, but you will find 
your own improvised lecture in the newspapers, 
and the radio will broadcast that according to 

you botanical study is very important for daily 
life of the people. 

When you tell people living along the coast that 
the brown algae like Sargassum has in principle a 
method of reproduction analogous to that of 
human beings, a clever boy may ask you to tell 
the how and why of this and he hears something 
about sperms and egg cells. A school teacher 
living at 1 km distance from the coastal mangrove 
asks you after your lecture to repeat for him what 
mangrove forest is and where he can find it. 
Local authorities want to hear from you about 
the possibilities of agar industry. Don't fall into 
this pit. 


Is a botanist in tropical regions only explorer 
and educator? No; he himself learns very much 
from the simple illiterate kampong people. 
They know their varieties. Some of them know 
trees in the forest, only from making a cut into the 
bark. Why not learn that? Some people may tell 
you about plants to be used as medicines. You 
may find drugs which are useful but unknown to 
the academic doctors, Indonesian or foreign, and 
you may find a good basis for botanical education 
of students in medicine and pharmacy. 


Especially growing institutes need a lot of local 
support and local understanding. At the start in 
Pajakumbuh, we were without housing and often 
without our own transport for excursions and 
expeditions. The behaviour of a botanist looking 
everywhere in gardens, in paddy fields, and in 
forests for plants was badly understood by the 
population. What do you look for? What is the 
purpose of your work ? I was asked these ques- 
tions everywhere. I made it the subject of a pub- 
lic lecture on the first anniversary of the Faculty. 
After that popular lecture, I got spontaneous 
help from the head of the local civil service to 
build an emergency herbarium, a building of 
8x14 meters from which several thousand dupli- 
cates are already distributed to London (Br. 
Museum, Nat. History), Kew, Leiden, Amster- 
dam, Geneva, and Singapore. Officers of the 
forest service joined several of my expeditions. 
The trip to the summit of Mt. Kerinchi was simul- 
taneously done by the head of Forestry with 
several of his officers. Many problems of forest 
botany, reforestation, and rural reconstruction 
are often discussed with my forestry friends, 
semi-academic Indonesians. All projects for 



nature protection are jointly made with the forest- 
ry service. 

The head of the service for agriculture is still 
like a father of the Faculty. He started it locally, 
and he supports our work in every way. He is 
as much disappointed as we when we cannot get 
the money and equipment from the Central 
government. With the head of the Veterinary 
service, we started our project for local recon- 
struction at the base of Mt. Sago. It is not neces- 
sary for us to tell the leaders of the Forestry 
service that their officers rarely enter the forests, 
to tell the agriculture service that its apparatus 
has very little practical use for the population, to 
tell the veterinary doctor that the grasslands on 
the experimental station are badly treated by 
burning, etc. They all know this, but they are 
waiting for our students who shall have to im- 
prove the present situation. The country waits 
for a new, better educated generation. We are 
clearly confronted with problems in our desa 
pertjobaan, rural reconstruction area, two kam- 
pongs with mixed gardens, surrounded by paddy 
fields, hills and with above them the forest of 
Mt. Sago. How are we able to advise, encourage, 
and educate the population here? Botani- 
cal inventories, socio-economic research by the 
students, practical projects for a cooperative 
movement including evening courses to be given 
by students are started here. Thus our work 
finds roots before modern laboratories and equip- 
ment are provided. Of course the illiterate popu- 
lation is not awakened within one year from its 
apathy, but there is more movement than we 
anticipated a year ago when this project was 


When agriculture students after three years of 
study receive only laboratory courses in botany, 
not any practical education in physics, chemistry 
and agriculture we can speak about mismanage- 
ment among the authorities who started this 
Faculty and who have the responsibility for its 
welfare. We must realize, however, that Indo- 
nesia is a land in transition with many financial 
and economic troubles. As long as these troubles 
are not resolved, reconstruction of the country 
remains difficult. It is, of course, disappointing 
to educate students in agriculture in botanical 
subjects while other subjects like chemistry and 
agriculture itself are largely postponed by lack of 
teachers, equipment, and working rooms. It is 
very irritating that improvement of this unsatis- 
factory situation is so long postponed. Visits of 


several foreign agencies UNESCO, Ford Foun- 
dation, 1CA, always came at the most inapprop- 
riate time. Now only some local oil companies are 
giving support (for chemistry). The best help we 
received was for the library, especially from Ame- 
rica, after the Botanical & Gardens of Bogor 
gave it a good start. 

As to botanical work, it is, of course, almost 
impossible to collect at new institutes within some 
years all the old books and journals which are 
necessary for revision work and monographs. 
This is another reason to concentrate on explora- 
tion, documentation in the field, and to distribute 
as many duplicates as possible of the valuable 
collections. The returning namings will help in 
the work of ecology and plant geography. 

The amount of work to be done (Stencilled 
lecture notes must be prepared for the students as 
no books in their own language are available, 
and they cannot read the Dutch language), is so 
over-whelming that almost no time is left for 
preparing scientific papers. Work of years ago 
still remains unpublished. Then we get requests 
from abroad to send material living or in liquid. 
We have to go to Java to get the chemicals for 
fixation; we are not able to obtain glasswork 
with stops and often cannot fulfill the wishes of 
foreign colleagues because of lack of time and 
provisions. The British Museum (Natural 
History) got a living specimen of Rafflesia by 
aeroplane. Nobody understands how many 
bureaucratic barriers had to be broken down for 
that within some days. I had first to get all kinds 
of permits from Djakarta and, was not allowed to 
pay until a Dutch firm took over the responsibil- 
ity. The Faculty waited two years before the first 
foreign journals entered the library. A large 
botanical institution in America, formerly special- 
ising in the flora of Malaysia, writes sour and 
formal letters to me in reply to enthusiastic pro- 
posals which are for mutual benefit. Other 
institutes like Amsterdam, Leiden, Cambridge, 
British Museum and Singapore are largely 
cooperating and stimulating. 


The work of botanical exploration and educa- 
tion in tropical regions needs the help and support 
from temperate regions of the globe. There are 
the books, the old publications, the old centres of 
botany, the treasures gathered by Blume, Miquel, 
Hooker, Ridley, De Candolles, and Merrill. 
There are still many more botanists working than 
here. Here we have the richest part of flora of 
the world still to be studied. 



What will be the future of the tropical forests? 
Will they be destroyed before scientific interest is 
raised among the people of these countries? 
Must all foreign botanists go away from here? 
Will it be possible for a botanist from Europe 
or America to devote his whole life to the richest 
flora of the world, or will political boundaries and 
nationalistic movements force him to restrict 
himself to the scanty flora in his own country or 
to shift his attention to parts of the tropics which 
are still under colonical rule? History teaches 
us that the fate and function of botany in our 
world cannot be separated from international 

In my opinion, this is understood by the grow- 
ing generation of Indonesian scientists, the best 

of them being further educated in Europe and 
America. The joint use of natural resources of 
the world will have its natural equivalent in joint 
exploration and research. 


This lecture started like the talk of a circus- 
artist and ended like a sermon. I hope that you 
understood that botanical exploration and educa- 
tion has found a good start in Central Sumatra 
itself, but that this young plant has still many 
difficulties in growth. The climate is often frus- 
trating and disappointing for it. Much skill, 
perseverance, and a sense of humour are needed 
for its growth. Have we to laugh about this 
undertaking or to pray for it? 





Kew Gardens, Surrey, England. 

In the modern world we are all increasingly 
dependent on the applications of science, and 
education for the young must include an intro- 
duction to both physical and biological science 
if they arc to have an intelligent understanding 
of their environment. And it is not only the 
young who need to have access to such knowl- 
edge; there is need for popular publications for 
adults, to keep them up to date with the changing 

All life is dependent on the green plant, and 
though modern technology has greatly changed 
our ways of living, it has not altered that fact. 
But the modern habit of living in towns and cities 
tends to prevent a great many people from 
realizing the fact, and therefore education in 
biology is of even greater importance than it was 
when the majority of people lived in close depend- 
ence on their natural surroundings. 

Modern political developments have added a 
new urgency to all aspects of education. Demo- 
cratic self-government, to be effective, depends on 
an educated electorate. 

Thus the first essential of education in biology 
is an understanding of the significance of plants 
as the basis of all life and as a basic natural 
heritage at the disposal of mankind, and education 
of this nature must be presented in terms of local 
plants in their relation to the local climatic 
environment. Knowledge of this kind is being 
gradually acquired at the various tropical bota- 
nical institutions. Such institutions thus have a 
duty to contribute knowledge about local plants 
which may be used in this necessary education in 
biology. And such institutions will in the future 
depend for their very existence on a general 
understanding of their importance by the public 
and by the politicians elected to power by the 
public. Publications of this kind become neces- 
sary publicity from the point of view of the 
institutions themselves. 

Biological science is not like physical science, 
the important data of which are independent of 
environment. Every different tropical country 
has different plants and animals, and there are 
many differences of climate which control and 

f Presented by J.V. Santos. 

limit the activity of those plants and animals, 
and also of man in exploiting them for his own 
use. Thus there is need for different presentations 
of these subjects in different countries; one general 
text book, or a series of publications from a 
single source, will not meet the situation. 

The great difficulty in preparing such educa- 
tional and popular presentations of a biological 
nature lies in the extreme complexity of the sub- 
ject matter and also in the fact that there has been 
far less biological study in the tropics than in 
temperate regions; and studies made in tem- 
perate regions are only of limited help because 
they deal with a different set of organisms in a 
different environment. 

Before anyone can write a simple introduction 
to a subject for the help of beginners, he must 
himself be master of the subject, or at least have 
a wide enough knowledge to be able to select for 
the beginner those aspects of it which will be most 
helpful. In order to obtain the necessary wide 
outlook, one must spend a great deal of time in 
study; and this involves not merely the study 
of books already written, but also the local 
plants themselves, as existing works cover only a 
fraction of what one needs to know. Original 
observation and the correlation of such observa- 
tion are creative processes quite different from the 
assimilation of knowledge already recorded in 
books. The ability to carry out such creative 
work is something that comes only from long 

As an example which has come to my personal 
attention, I will refer to the question of bamboos 
in Malaya. Bamboos are of great importance to 
the countryman who uses them every day in 
many ways, and any student of biology in Malaya 
should learn something about them. But there 
is nothing in print about bamboos in Malaya 
which is of any use to a person who wishes to 
know how to begin to study the subject, nor 
even any work which is of direct value to the 
expert; the only usable taxonomic works have 
been written in India and in Java, and they leave 
untouched much of what is in MaUiya. Further- 
more, nobody has given critical thought to the 



general classification of this group of plants as 
a whole (quite apart from Malaya) for nearly 
a century. Some twenty years ago I began to 
take an interest in the subject, and after a good 
deal of casual enquiry I sat down to write a sys- 
tematic account of what information was avail- 
able, based on specimens and data accumulated 
in Singapore. This showed me how inadequate 
this information was, and during subsequent 
years I made several journeys with the object of 
adding to it. Having left Malaya, I cannot add 
further original observations, and I have written 
an account of the subject, as far as is possible on 
present information, so that others may start 
where I leave off; in particular, I have tried to 
present an introductory statement which will 
help a new observer to begin his work. 

This work on bamboos is only a small aspect 
of the study of Malayan plants, and it may be 
matched in other families of plants and in other 
parts of the tropics. But unless this kind of work 
is done, the peoples of the world will not be able 
to know the scope of their natural heritage of 
plant life, and they may well, in ignorance, destroy 
a large part of that heritage before it has been 
studied. Such primary study of native plants 
can be accomplished only by experienced workers ; 
it is not something that can be undertaken by a 
beginner. This is often not understood by ad- 
ministrators and politicians, and even scientists 
who are not acquainted with the problems of 
taxonomy sometimes fail to appreciate the great 
amount of such work which remains to be done 
in the tropics. 

Of course there is a great amount of informa- 
tion already recorded about tropical plants. 
Much of this information is in taxonomic mono- 
graphs and formal Floras which can be under- 

stood only by a specialist. There is need, as above 
noted, for specialists to select from this mass of 
data such parts as are useful as an introduction 
to the subject. The university student, or the 
school teacher in training, needs to have such an 
introductory presentation. When the student 
in turn becomes a teacher in a school, he will 
find yet another problem that of introducing 
children to the same kind of knowledge. The 
teachers in their turn must thus review the subject 
and write books which will serve their particular 

From my present standpoint, the work of the 
tropical botanical institution is, therefore, two- 
fold: the continued study of natural vegetation 
and the recording of the results of that study in 
formal scientific publications and the interpre- 
tation of that knowledge in terms useful to the 
student (especially the student who will be a 
teacher), and to the adult who wishes to develop 
an interest in such matters. 

The institutions will be able to do tins only if 
there is a sufficient reserve of natural vegetation 
and if they have sufficient trained and experienced 
stalT to carry out both the fundamental investi- 
gation and its interpretation for the student and 
the common man. In these days, the institutions 
will not be provided with the staff, equipment, 
and means of publication unless the general 
public is aware of the importance of these things. 
The right kind of publicity is needed, and this 
can come only from the institutions themselves. 
It should be a function of this Congress to help 
to impress on governments and on peoples that 
the work of tropical botanical institutions, and 
with it the preservation of natural reserves of 
plant life, is a matter of fundamental and urgent 
practical importance. 


j.v. SANTOS: Mr. Holttum appears to support Van 
Steenis' idea. 

C.G.G.J. VAN SIEENIS: Dr. Holttum in his paper in- 
tended that there should be more simple, popular knowl- 

edge of plants. Dr. Holttum has written papers not 
intended for secondary schools. Structure must start first. 
F.R. FOSBERG: 1 am pleased to hear this discussion 
about starting education in secondary school. 





Department of Botany, University of Hawaii, Honolulu 14, Hawaii. 

The algal roles in nature at sea are those of 
being the primary producers: primary producers 
of food and materials in the euphotic zone. The 
rates of algal production and perhaps the sizes 
of algal standing crops are a function of the con- 
centrations and rates of addition of otherwise 
limiting chemicals to this euphotic zone and to 
light intensity. To an extent these quantitative 
population features are a function of the abilities 
of the organisms themselves. The major roles of 
the algae are related to whether they are plank- 
tonic or benthic in nature. 

In the open sea the algae are plankters, which 
in at least some cases must lead a heterotrophic 
life. As an example, the Coccolithophores domi- 
nant in euphotic Mediterranean waters (3) have 
also been found to be abundant at depths of 1,000 
to 2,500 meters, far below the euphotic zone. 
Such organisms may get there initially through 
sinking or by vertical mixing. In our own work, 
we have upon occasion found the most chloro- 
phyll-rich waters at the very bottom of the eupho- 
tic zone, if not below it. Returned to the surface 
through upwelling or vertical mixing or rising 
through changes in density, such organisms may 
act as the "seed" for normal populations. 

The food producers in the sea, the phyto- 
plankton algae, make that 73 per cent of the 
earth's surface covered by ocean as productive as 
the land on the average per unit of area. This 
production may be estimated variously (e.g., 20, 
22, 9) between 0.8 and 15 x 10 l tons of carbon 
per year. 

The rate of primary production varies in dif- 
ferent parts of the ocean. At the poleward limits 
of the temperate seas with seasonal change and 
the vertical mixing of various kinds that occurs 
there, the substances normally limiting to phyto- 
plankton production become more readily avail- 
able. Measurements of phytoplankton product- 
ivity have been found (15) to increase three-fold 
as one moved into this productive water from the 
temperate region. In the divergent current 
regions of the Central Pacific along the equator, 

water from the dysphotic zone upwells. In such 
regions the productivity has been measured (16) 
to be almost ten times that several degrees to the 
north or south. In these two instances, the rate 
increases are correlated with areas where the 
inorganic materials normally limiting to algal 
growth are being brought upwards into the eupho- 
tic zone from below. These areas of major size 
and consistent presence may be correlated with 
high fish production: albacore tuna in the first 
instance cited and yellow-fin tuna in the second. 

Phytoplankton productivity rates increase as 
land is approached. This has been shown (12) 
to be an increase of two or more orders of mag- 
nitude. The phenomenon is apparently due to 
two processes leading to high concentrations of 
essential minerals near shore. They are (a) con- 
tributions from the land brought by water, and 
(b) substances accumulated in the area from the 
sea by benthic forms. 

General algal productivity in the non-enriched 
areas of the sea may be supported by recycling of 
materials derived from the sea or initially from 
emergent land. Such areas may gradually become 
poorer in the elements essential for primary 
productivity through the movement of materials 
to layers of water below the euphotic zone. 

Fixed nitrogen may be exceptional in that 
supplies of this element in combined form may 
be derived from rainfall or may dissolve in the sea 
and be combined there by microorganisms, 
particularly the blue-green algae. In fact on 
atolls and in the sea, nitrogen fixation by blue- 
green algae is expected to be a major as yet non- 
assessed factor. If the major premises of this 
and the previous paragraphs are valid, then a 
relative increase in the ratio of nitrogen to, e.g., 
phosphorus might be measured in going away 
from areas where there is enrichment from shore 
or the depths. 

It is of considerable interest to note the trend, 
even in the study of benthic algae, away from 
more descriptive aspects and toward more func- 
tional aspects. Thus in past years we had oceanic 

t The conclusions presented herein without citation were drawn from studies financed largely by contracts between the 
Botany Department of the University of Hawaii and the U.S. Atomic Energy Commission (AT-(04-3)-15) or the U.S. 
National Science Foundation (NSF G 1992 and NSF G 3833) or with the U.S. Office of Naval Research (NR 388-001) 
with the U.S. National Research Council's Pacific Science Board, and by funds from the Graduate Research Committee 
of the University of Hawaii. 




phytoplankton studies concentrating on observing 
the kinds and numbers of algae. The outstanding 
example of this is the magnificent series of studies 
by Lohman (17). Today we have concentration 
on studies of oceanic phytoplankton such as 
those of Steemann-Nielsen and Aabye-Jensen 
(23), using isotopic carbon, and those of Ar- 
rhenius (this Congress) on bottom deposits where- 
in only the algal function or results of algal 
functions are observed and measured. But very 
few, an example being the fresh-water lake studies 
of Rodhe (21) have been able to link both obser- 
vational and functional studies. Marine algal 
studies of this kind should be encouraged. 

The benthic algae are more familiar and are 
the forms thought of as the marine algae first, 
even by phycologists themselves. These are the 
algae normally living attached to the substratum 
and commonly ranging in size and form from 
half-ton kelps to solitary diatom cells. In com- 
parison to the plankton algae they are very well 
known taxonomically and morphologically. For 
reasons of economic interest, species of some 
genera, such as Chondrus, Macrocystis, Laminaria, 
Gelidiwn, Eucheuma, Gratilaria, and Porphyra 
are rather well known, even in regards to their 
production (e.g., 28 on Gelidiuni). Their con- 
tributions otherwise are conjectured but little 
assayed as yet. Two roles can be expanded upon 
here as conspicuous in the more central Pacific: 
these are essential mineral concentration near 
shores and material construction of shores and 

The mangrove areas in the Hawaiian Islands 
are conspicuously clean and devoid of the char- 
acteristic faunal and floral populants. In the few 
years mangroves have been planted there, the well 
known algal combination of Caloglossa, Bostry- 
chia, and Catenella apparently has not appeared. 
Surprisingly enough some of the mangrove areas 
studies on Mindoro (east of Calapan), the Philip- 
pines, and in the Thousand Islands (northwest 
of Djakarta), Indonesia, were similarly devoid of 
these macroscopic algae. But then, too, these 
localities were devoid of the intertidal mud char- 
acteristic of the areas heavily populated (e.g., 
Pine River shores in Queensland, Australia) by 
the above algal triumverate. The theory of 
causality rears its head at this place: mangroves 
develop on reef flats and elsewhere where no mud 
is present, but do the algae appear where there 
is mud, or, do they cause the mud to be deposited ? 
In the Pandan Nature Reserve in Singapore 
where there is a magnificent development of 
these algae on both mud and mangrove roots, 

the algae were seen to be muddy, covered with 
a seemingly inseparable coating of mud, while 
the roots on which they were growing were, 
otherwise, quite devoid of any regular deposit 
of mud. In fact the roots except for the algae 
were as clean as those in Hawaii or the roots of 
"non-algalated" mangroves in Indonesia or the 

In Hawaii, in Singapore, and elsewhere, 
Vtntcheria forms dense turfs of erect filaments on 
flat mud areas of deposit, at the edges of drainage 
channels. It appears, from their mud-choked 
flat nature, that these turfs act as mud-trapping 
agents. The presence of this phenomenon far 
outside the mangrove area (e.g., at Coos Bay, 
in Oregon, U.S.A. 5) inclines one to exclude this 
community from the mangrove community, 
though it is strongly developed in some mangrove 

Reefs and non-igneous islands as we know 
them in the Pacific are of biological origin and 
largely rather directly the results of algal activity. 
Reefs as we are using the term are solid aggre- 
gations largely of carbonate, which has usually 
been deposited by Rhodophyta, Foraminifera, 
Coelentcrata, or Chlorophyta. The order of 
abundance in a reef is often listed in order as 
these groups appear above. 

This definition of reefs excludes wave cut 
benches and solution benches, though it is postu- 
lated here that some of their common physiog- 
nomy is caused by the same effects of algal 
growth. Both the benches and reefs often have 
an outer raised margin (rampart or algal ridge) 
and a higher algae-covered area of reef flat just 
inshore of the outer margin. Wentworth (26) 
attributes this form in the case of the benches to 
an algal function: that of reducing erosion. On 
reef edges such as found in the Tuamotu Archi- 
pelago (11) in addition to building the reef margin 
upwards, the algae seem to play the same pro- 
tective role seaward of the channel area (back- 
ridge trough). Thus a common role of the algae, 
mitigating the results of erosion, is postulated 
here as a major cause of this similarity in form 
often seen on all three shore types. 

In different places and perhaps at different 
geological times, the relative importance of the 
above groups of organisms giving rise to coral 
has changed. As an example it may very well be 
that among Chlorophyta, formerly the Dasycla- 
dales were more important than now. In atolls 
seaward reef edges are usually predominantly 
red algal (Corallinaceae) in composition while 



those facing lagoons are predominantly coelen- 
terate in composition. Paleontological study of 
reefs has been of academic interest until recently 
when certain ore and oil associations with fossil 
reefs have brought economic reasons to bear on 
a better understanding of reefs. 

Coral islands are common in the Western 
Atlantic and elsewhere. Here this term is used to 
denote islands the detrital material of which is of 
biological origin but islands which are not based 
on or surrounded by a currently active reef flat. 
Perhaps those low islands, such as the Bahamas 
in the Caribbean or the Thousand Islands in In- 
donesia, are of this general sort. Often they are 
largely of algal detritus. 

The coral islands arc of several types including 
those which are perhaps relics of the normal 
reef island situation to be discussed below. John- 
ston and Sand Island at what may be called here 
Johnston Atoll (south of Hawaii) and some of 
the smaller islands of the Hawaiian group (e.g., 
Laysan) are perhaps of this sort. The composition 
is generally thought of as predominantly algal 
but is often line or for other reasons hard to assay 
as to origin. Some of the Bahama group and 
Bermuda have been described as consisting largely 
of Halimeda detritus. These are of easily recog- 
nizable rather uniform fragmental composition. 

Coral islands of animal skeletons, in addition 
to the two algal types above, are found. The 
Thousand Islands near Djakarta seem to be at 
least in part of this sort. It may be that such 
islands arc more frequent in the Eastern Indo- 
Pacilic and, having been investigated first, gave 
rise to the seemingly false idea of the dominance 
of coelenterate corals in reefs and islands of bio- 
logical origin. Such islands, as typified by those 
near Djakarta, when small, slope gently from at 
least ten meter depths (as judged by gradual shift 
in water color) to low tide level and, then, termi- 
nate upwards in low (one meter above high tide 
line) Hat island fragments. Larger islands may 
have reef flats that appear to slope somewhat 
toward the island. In such cases mangroves, 
instead of covering the intertidal shore or reef 
flat, are at times separated from shore by a body 
of water, which might be termed a lagoon. If 
one terms the shallowest water at the seaward 
margin of such a lagoon, a reef, it must be kept 
in mind that it is different from the reefs domi- 
nated by red algae in at least two respects aside 
from the actual organisms present. First, such 
reefs seem to be gently rounded in contour, in 
their shallowest areas or slope steadily into the 
sea rather than being quite plane or even with a 
raised margin as in the case of algal reefs. 


Secondly, it appears that they do not extend 
upward to such a level that they are exposed to 
the air as much as are the red algal reefs. It 
appears that a majority of Central Philippine 
shore reefs are of this latter type. 

Though structurally of coelenterates, this last 
type of coral island, or reef, is inseparable from 
the zooxanthellae and filamentous algae which 
pervade it. Though far from satisfactorily under- 
stood, it appears that hermotypic corals, the large 
and only corals contributing significantly to reef 
and island substance, are dependent upon these 
algae. It may be debated whether this dependence 
is nutritional or functions more in removal of 
waste products. Recent work (1, 13) showing 
the loss of a large part of the photosynthate from 
unicells would lead us to expect the nutrient was 
available, in addition to the older arguments that 
these endophytes were nutritive; e.g., hermotypic 
corals develop only in the euphotic region. Un- 
doubtedly the algae untilize the mineral waste 
products that diffuse away from the coelenterate 
cells, but it seems less likely that this is essential 
to the livelihood of the animals. If the hermo- 
typic coelenterates are dependent on the algae, 
then in turn we can say that even the animal 
dominated reef structures are quite directly related 
to algal activity. 

A note at this point is to be entered as to the 
effects of algae on coral island (19) intertidally 
and above high tide line. Their growth on almost 
all the surfaces of these low oceanic islands is 
a phenomenon associated with the marine en- 
vironment. The gastropods in rasping them away 
as food remove consolidated materials. Their 
growth on, and in, coral rock must effect its 
solution. These two phenomena are blue-green 
algal phenomena and as the major of these 
algae, Entophysalis, is also abundant on much of 
the non-regularly inundated part of the island, 
the general phenomenon of nitrogen fixation by 
such algae should not be overlooked. On these 
low oceanic islands, the usual nitrogen fixing 
organisms, such as the Rhizobium of legume 
roots, are essentially absent. In wetter areas, 
such as Majuro in the Southern Marshall Islands, 
the island surfaces, where devoid of larger vege- 
tation, may become covered by Nostoc commune. 
Such algae have been shown (27) to fix up to 90 
per cent of the nitrogen they use even when sup- 
plied with what would appear to be adequate 
supplies of fixed nitrogen. 

Coral reefs as found in the Central Pacific and 
the islands on their reef flats seem to be main- 
tained at the sea surface by algal activity. Char- 
acteristically, atolls are formed. There is, as 



often presented, a variety of reefs in relation to 
igneous islands or underwater bases. The theore- 
tical origins of atolls and the accumulation of 
islands on atoll reefs have been elaborated on at 
length by many authors. Other than to note 
that these authors have concerned themselves 
with the formation of "the classical atoll in full 
bloom" from its beginnings and have given little 
attention to what may be called their old age or 
decline, we will restrict ourselves to the postulated 
algal roles not discussed above. 

Let us first consider in brief the distribution of 
algae across an atoll. As an example let us con- 
sider Raroia in the Eastern Tuamotus(7,//,/#J or 
Eniwetok in the Northern Marshall Islands 
(25, 9) i as the algae can be found in an investi- 
gation leading in over the upwind seaward reef, 
moving downwind, across the lagoon and thence 
traversing the leeward atoll rim. Upon approach- 
ing the atoll coming downwind, the water 
becomes shallower as the bottom slopes gently 
toward the surface. Rather abruptly at depths 
of about six to eight meters, the hermotypic corals 
become covered or arc replaced by coralline 
algae. In shallow water, there is little else in 
sight. In deeper water, coclenterate corals arc 
scattered irregularly over the gently undulate 
surface radiating away from the atoll. In shallower 
water, radiating rows of individual coelenterate 
coral organisms give way inshore to grooves and 
spurs covered with a smooth coating of algal 

Spurs extending toward the atoll, at the sea 
surface, may extend upwards to such levels that 
the seaward margin of the reef protrudes perhaps 
a meter between waves even at high tide. The 
spurs tend to be especially well developed oppo- 
site islands and may be fused laterally into a 
ridge, the famous algal ridge of the literature. 

Grooves extending toward the atoll may run 
through the algal ridge or terminate outside of it. 
They may extend through the ridge covered by 
the laterally fused ridge material and reef surface 
inshore of the algal ridge. They may be open 
channels throughout or closed over so that the 
innermost open end is a blow-hole, sometimes 
enclosed in a low dome, on the inner part of 
the reef. 

Inshore of the algal ridge, the surface is usually 
plane, horizontal and at low tide level. This 
surface may be as smooth as a concrete sidewalk. 
Sometimes it is covered with a turf of jointed 
coralline or other algae. As Wentworth sug- 

gested (26) for solution and wave-cut benches the 
algae here also probably play a role in lowering 
erosion, as discussed above. 

Still further inshore, the reef surface becomes 
lower and tends to slope so that water coming 
in over the algal ridge flows to the right or left 
and back out over the reef margin through 
grooves or the so-called (11) excurrent areas. 
This area has been termed a back-ridgc-trough 
and termed elsewhere a channel area. This area 
appears to be one where corrosion and corrasion 
are dominant over deposition. Indeed from the 
algal pavement of the outer reef flat to the lagoon, 
the physical features appear to be largely regulated 
by mechanical forces though the materials may 
be predominantly of algal origin, and these latter- 
may in turn regulate the mechanical forces. 

The sea reef, phycologically, can be divided 
into four major zones horizontally in so far as 
the populations are concerned: (a) Porolithon 
onkodes dominates the surfaces from depths of 
six to eight meters up onto the outer reef flat; 
(b) the outer and higher reef flats which may be 
covered by an algal turf; (c) the channel or reef- 
pool area; and (d) the shore intertidal and supra- 
tidal areas. In designating these we purposely 
avoid terms used in discussions of vertical dis- 

Porolithon onkodes above its maximum depth 
comes quickly to cover almost all of the surface. 
It produces the algal ridge as a scries of overlaying 
overlapping pustules of several centimeters to 
a few decimeters in lateral extent and from a few 
millimeters to a centimeter or two thick. Within 
the cavities resulting reside a host of animal sorts, 
of which annelids, fishes, crabs, and foraminifera 
are conspicuous. On the surface patches of small 
algae, e.g. Centroceras, Microdictyon, and blue- 
greens, are scattered about as are such animals 
as limpets, barnacles, and Cypraeids. Poroli- 
thon onkodes is a most peculiar coralline alga in 
that it will live on the most brilliantly illumi- 
nated shore areas and stand some exposure to 
air. This is in contrast to other genera of crustose 
coralline algae such as Lithothamnion^ which 
never grows extensively under such circumstances. 

The algal turf on higher parts of the reef flat 
occurs in at least two major forms: as relatively 
pure stands of jointed corallines on outer reef 
flats (7) underlain by Porolithon onkodes ; and as 
mixed stands, on inner reef flats (24), of Jan fa. 
Amphiroa, and relatively small fleshy algae such 
as Caulerpa ambigua, Polysiphonia, Herposiphonia 

1 Perhaps the confusion regarding Lithothamnion and the algal ridge arises from the fact that Porolithon onkodes was 
originally described by Heydrich in the genus Lithothamnion. 



secunda, Laurencia, and Gelidium intermixed often 
with large proportions of blue-green algae. Larger 
fleshy algae often occur in patches or isolated 
clumps or sandwiched in crevices or the holes 
opened by boring animals, i.e., where freed from 
the browsing of fishes. Two functions must be 
considered for this turf: first, in providing a direct 
or indirect source offish food; second, in reducing 
erosion of the reef flat. 

The channel or pool area is often sandy or 
floored by shallow pot holes. Algae are few 
except under the edges of reef boulders, e.g., 
Diciyosphaeria. Often the surface is pinkish 
from what seems to be incipient growths of 
PoroUthon onkodcs. Larger algae, such as Liagora, 
Tur binaries and Padina sometimes occur. The 
aspect is that of a shallow but open tidepool. 
At one time (February), the reef pool area of the 
eastern end of Ujcland in the Marshalls was 
strongly pink from the air. Investigation on the 
ground revealed this to be due to an extensive 
growth of Liagora. 

The intertidal and supratidal areas are usually 
covered by, below, a brown and, above, sun- 
blackened coatings of Entophysalis. As one pro- 
gresses away from shore and salt, Anacystis and 
finally Scytonema or, if very wet, Nostoc similarly 
tends to cover stable surfaces. The roles of these 
algae in nitrogen fixation and shore processes are 
taken up above. 

Upwind lagoon shores are usually sloping sand 
beaches. There is little of macroscopic biological 
nature on such sandy beaches. When not sandy 
beaches, they are representative of shore types 
elsewhere in the lagoon. The color of the water 
changes as one moves out into the lagoon from 
the shore. As seen from the air, the color is that 
of the sand beneath, usually a very light buff 
color, darkening until depths of six to eight feet 
arc reached when the color is definitely green 
tinged. The water appears greenish and gradually 
darkening until at perhaps depths of twenty feet 
when it can be said to be blue. When shallower 
places in the lagoon are approached, a reverse 
color sequence shows. 

The most shallow places in lagoons are the 
brown, sometimes pink-edged or light buff- 
centered reef patches, reaching low tide level. 
Chunks of rock protruding above low tide level 
are often blackened with blue-green algal growths. 
If islands are present, they are likely to be similar 
to the sandier reef islands. Reef patches have 
various forms (4) which may be arranged in an 
evolutionary sequence (ll)\ either in the lagoon 
changing in relation to the bottom, or just growing 


upwards, or in relation to changes in the shor 
location with time. 

A typical reef patch reaching the low tide sui 
face of the lagoon is coated on its sides wit 
Caulerpa and, especially on less precipitous area 
and near the base, with Halimeda. The Caulerp 
is reputed by the Tuamotuan natives to be a mos 
important turtle food. Small fleshy algae occui 
especially on the uppermost six to eight feet c 
the rocky column, and serve as fish food. Th 
classical relationship between the coelenterate 
and the algae perhaps enables the hermotypi 
corals to grow and play their role in the produc 
tion of the reef patches. The brown coloratio 
of the coelenterate corals on top of the reef pate 
is probably largely due to the endozoic algs 
pigments. Occasionally the margins of rcc 
patches will be pink from growths of PoroUthon 
especially on their upwind sides, or they ma 
bear a fur of Rhodomelaceous algae. The center 
of reef patches are often depressed and san< 
floored with scattered coral fragments. Ther 
above the sand such algae as Udotea and th 
turf-forming species of shore areas are oftei 

Diving and careful dredging in lagoons hav 
contributed (9J4) considerably to our knowledg 
of the algae on lagoon bottoms. With the ex 
ception of Halimeda, little in the way of corallini 
or calcium-depositing algae is found. Contrar 
to popular opinion, Halimeda has not been foun< 
( 11 J 4) to occur in dense meadows on the bottom 
Instead, though lagoon bottom sediments ar< 
often predominantly of Halimeda segments, th 
living thalli are almost always clustered abou 
the talus bases or occur on the sides of ree 
patches. Despite this, Halimeda is perhaps th< 
agent most responsible for the filling of lagoons. 

Reefs are usually found on the downwinc 
lagoon shores. These have been termed f/7 
shore-bound reef patches for both their anatonr 
and components are very similar to what is founc 
on the lagoon reef patches. The roles of the alga< 
seem to be the same as on lagoon reef patches. 

Islands on the downwind reefs are, phycologi 
cally, very similar to those on windward reefs 
The islands which occasionally occur on ree 
patches are also phycologically similar. 

Seemingly downwind or lea sea reefs are phy 
cologically like those of the windward shores anc 
the algal roles are the same. In some cases, e.g. 
at Arno Atoll in the Southern Marshall Islands 
the downwind reef edge may have many mon 
coelenterate corals on it than the sea reef edges 



elsewhere, and also the algal ridges are often less 

As described above, the atoll system seems to 
persist in equilibrium with the sea as long as 
changes in the earth's crust are slow. If the crust 
shifts upward, the reef can be eroded back toward 
sea level; or if the rate of elevation is sufficiently 
fast, a raised atoll can result. On the other hand, 
should the atoll sink, upward growth of the 
organisms would keep the atoll in contact with 
the sea's surface, unless the sinking rate were 
so fast the organisms could not do so. Apparent- 
ly a sinking rate of over one or two centimeters 
per year would result in a sunken atoll, at least 
that is the current estimate for the maximum rate 
of atoll upgrowth. Examples of atolls where 
such phenomena have taken place arc numerous. 
The atoll around Johnston Island, south of 
Hawaii, seems to have sunken on only one side; 
so a long crescent rim is left at the sea's surface. 
The atoll of Anaa, east of Tahiti, is an example of 
an atoll where one edge has been raised, in this 
case about five meters, while the rest is at normal 
atoll elevations in respect to the sea surface. 
Prospective algal roles in both the erosion and up- 
building have been taken up above. 

Finally the benthic algae seem to play a most 
important role in the concentration of the che- 
mical elements essential to life processes. As a 
result of this process, as one nears shores the 
biological standing crops as well as the productiv- 
ity rates increase enormously. Let us consider 
the events that would lead to this end around a 
newly created igneous island such as a volcano 
suddenly rising above the sea's surface. Essen- 
tially this condition prevails in Hawaii when new 
lava flows reach the sea. From a study there of 
lava surfaces appearing in the sea in 1955 we know 
it is the algae which appear first and before the 
animal populations characteristic of older shores 
in the same general area. 

In the open ocean we would assume that when 
a new igneous mass appeared there would at first 
be no concentrations around its shores. Leached 
materials from the newly cooled lava would be 
diluted and washed away in the passing ocean 
stream. The first macroscopic organisms to 
appear would be expected to be such as Enter -o- 
morpha, Ectocarpus, and Polysiphonia, as they are 
in Hawaii, if the new island were in similar waters. 
These pioneers appear in that order of importance 
and time in the Central Pacific. Other populants 
follow. The benthic standing crop increases as 
the algae convert inorganic materials from the 
water into organic materials or deposit them as 

inorganic compounds such as calcium carbonate. 
This must apply as well to phosphorus and 
nitrogen and carbon and the other elements 
essential to protoplasm. Because these popula- 
tions are benthic forms they do not wash away. 

Following the algae, there appear the animals 
which feed upon the algae: the littorines, limpets, 
and blennies. A certain amount of the increase 
in standing crop shifts over to these herbivorous 
forms and then, theoretically, to carnivores. The 
detritus and fecal material are undoubtedly de- 
graded to a mineral state, and thus the water 
near shore becomes enriched in inorganic nu- 
trients. The observed near-shore increases in 
plankton must be the results of this enrichment. 

It is observed that the calcareous organisms 
appear conspicuously a year or more after the 
non-calcareous algae arc well established. With 
the very long-term evolution of an atoll, the 
binding of materials becomes predominantly an 
activity of the calcareous forms in so far as mass 
is concerned. This implies a shift to such herbi- 
vores as the parrot fish, some species of which 
live on calcareous algae. The detritus and fecal 
material they produce contribute to the sediments 
of the lagoon or island shore as well as to the 
enrichment of the waters about the high island 
or atoll. 

It is expected that the supply of fertilizer 
elements and the standing crop of organic material 
would increase until a new steady state is achieved. 
Previous to this time, i.e., while the standing 
crops are increasing, one might expect the water 
approaching the young atoll to contain more of 
the fertilizer elements than the water leaving the 
atoll. This is without taking into account sub- 
stances from a high island around which the atoll 
might be developing. In later stages, the amount 
would be expected to be equal. Perhaps in old 
age the amount leaving (at least leaving the 
euphotic zone) would exceed the amounts arriving 
and being bound by the primary productivity of 
the algae. It may be that plankton algal pro- 
ductivity near shores, as well as the size of the 
standing crops of phytoplankton, may serve as 
an index of the relative age of an atoll. 


The phytoplankton algae are described as 
having major roles affecting fertilizer concentra- 
tions or in primary productivity resulting from 
variations in fertilizer salt concentrations found 
(a) in transition or shear zones enriched by ver- 
tical mixing or eddies, (b) in divergent current 



areas by upwelling, (c) in wind disturbed areas 
by vertical mixing, and (d) near land by fertilizer 
increases arising both through concentration 
by benthic organisms and through addition by 
fresh-water. The benthic algae are described as 
having major roles in primary productivity of 
food, in the fertility of the soil, and in construction 
and destruction of shores, islands, and reefs, 
particularly those of atolls. 


(1) Allen, M.B., 1956, Excretion of Organic 

Compounds by Chlamydomonas, Arkiv 
F. Mikrobiol., 24: 163-168. 

(2) Arrhenius, Gustav, (Mss.) Climatic Records 

on the Ocean Floor. (Paper presented 
to symposium on marine upwelling at 
9th Pacific Science Congress, Bangkok, 

(3) Bernard, F., (Mss.) Donnecs R6centes sur la 

Fertilite Elementaire en Mediterrance. 
(Paper presented to the 1957 I.E.S. 
annual meeting in Bergen, Norway.) 

(4) Denielsson, Bengt., 1954, Native Topograph- 

ical Terms in Raroia, Tuamotus, Atoll 
Research Bulletin, 32: 92-96, 3 figures. 

(5) Doty, Maxwell, S., 1947, The Marine Algae 

of Oregon, I: Farlowiu, 3: 1-65. 

(6) , 1 954a,* Current Status of Carbon- 
fourteen Method of Assaying Productiv- 
ity of the Ocean (As of April, 1954), 
9 pp., 6 appendices, University of Hawaii. 

(7) , 1954b, Floristics and Plant Ecol- 
ogy of Raroia Atoll, Tuamotus, Atoll 
Research Bulletin, 33: 1-41, 1 figure. 

(8) , 1955,* Current Status of Carbon- 
fourteen Method of Assaying Productiv- 
ity of the Ocean (As of February, 
1955), 52 pp., 4 appendices, University 
of Hawaii. 

(9) , 1956,* Current Status of Carbon- 
fourtecn Method of Assaying Productiv- 
ity of the Ocean (As of April, 1956), 
51 pp., 6 appendices, University of 

(10) , 1957,* Current Status of Carbon- 

fourteen Method of Assaying Productiv- 
ity of the Ocean (As of July, 1957), 
41 pp., 2 appendices, University of 

(11) Doty, Maxwell S., and Morrison, J.P.E., 

1954, Interrelationships of the Organisms 
on Raroia aside from Man, Atoll Re- 
search Bulletin, 35: 1-61, 8 figures. 

(12) Doty, M.S., and Oguri, M., 1956, The Island 

Mass Effect, Journal du Conseil pour 
V Exploration de la Mer, 22: (1) 33-37. 

(13) Fogg, G.E., (Mss.) Extracellular Products of 

Phytoplankton and the Estimation of 
Primary Production. (Paper presented 
to the 1957 I.C.E.S. annual meeting in 
Bergen, Norway.) 

(14) Gilmartin, MalvernJr., 1956, The Ecological 

Distribution of Certain Central Pacific 
Atoll Benthic Algae, iii and 78 pp., 8 
ligures, and a map. M.S. Thesis, Univer- 
sity of Hawaii, Honolulu (Un- 

(15) Guillard, R.R.L., Doty, M.S., and Oguri, 

M., 1955, The Productivity of Waters 
North of Hawaii as Determined by 
Carbon-fourteen Uptake. Abstract from 
the Proceedings of the Hawaiian Acade- 
my of Science, 1955-1956, p. 10. 

(16) King, J.E., Austin, T.S., and Doty, M.S., 

1957, Preliminary Report on Expedition 
"Eastropic". U.S. Fish and Wildlife 
Service, Special Scientific Report 
Fisheries 201, 155 pp. 

(17) Lohmann, H., 1908, Untersuchungen zur 

Feststellung des vollstandigen Gehaltes 
des Meeres an Plankton, Komm. z. 
Wiss. Unters. D. Deutch. Meere in Kiel 
und d. Biol. Anst. Helgoland. Wiss. 
Meeresunters, N.F. Abt. Kiel. 10: 

(18) Newell, N.D., 1954, Reefs and Sedimentary 

Processes of Raroia, Atoll Research 
Bulletin, 36: 1-35. 

(19) Newhouse, Jan., 1954, Ecological and Flor- 

istic Notes on the Myxophyta of Raroia, 
Atoll Research Bulletin, 33:42-54, 2 

(20) Riley, Gordon, 1953, Letter to the Editor. 

Journal du Conseil pour r Exploration 
dela Mer, 19(1): 85-89. 

(21) Rodhe, Wilhelm, (Mss.) The Primary Pro- 

duction in Lakes: Some Results and 
Restriction of the C 14 Method. 

* These are annual reports to the U.S. Atomic Energy Commission^ Division of Biology and Medicine, of work accom- 
plished under contract AT-(04-3)-15 between the U.S. Atomic Energy Commission and the University of Hawaii. 
They have not been published and are not generally available but are listed here for convenience in completing the list 
of information sources. 




(22) Steemann-Nielsen, E., 1952, The Use of 

Radio-active Carbon (C 14 ) for Measur- 
ing Organic Production in the Sea, 
Journal du Council pour V Exploration de 
la Mer, 18(2): 1 17-140, 7 figures. 

(23) Steemann-Nielsen, E., and Aabye-Jensen, 

E., 1957, Primary Oceanic Production. 
The autotrophic production of organic 
matter in the oceans, Galathea Report, 
1:49-136, 41 figures. 

(24) Taylor, W.R., 1950, Plants of Bikini and 

other Northern Marshall Islands. Uni- 
versity of Michigan Studies, Scientific 
Series, 18, 15 and 227 pp. Plates 1-79 
and frontispieces. 

(25) Tracey, J.l.Jr., Ladd, H.S., and Hoflrneistcr, 

J.E., 1948, Reefs of Bikini, Marshall 
Islands, Journal of the Geological So- 
ciety of America, 59:861-878, Plates 
1-1 1 and 8 figures. 

(26) Wentworth, Chester, 1939, Marine Bench- 

forming Processes, II : Solution benching. 
Journal of Geomorphologv, 2: 3-25, 12 

(27) Williams, A.L:., and Burris, R.H., 1952, Ni- 

trogen Fixation by Blue-green Algae and 
Their Nitrogenous Composition, Ameri- 
can Journal of Botany, 39: 340-342. 

(28) Yanegawa, T., and Tanh, K., 1957, Studies 

on Agar-agar in Japan, Proc. Eighth Pac. 
Sci. Cong. (Philippines), 3: 215-223. 


R.C. MURPHY: Will the mere shallowing of water, as 
an ocean current approaches an island, cause a concen- 
tration of plankton life, and therefore increase the food 
resources for fish and birds? 

M.S. DOTY: In Hawaii, in respect to the circulation in 
the North Pacific, there is no upwelling along the islands. 

G.L. CLARKE: As the contained life is an integral part 
of the water volume, no such vertical concentration of 
plankton can occur. I attribute the acknowledged enrich- 
ment of life in the coastal waters to more favorable con- 
ditions for nutriment and reproduction. If the animals 
are brought by currents, they will also be carried away by 
currents that pass around the island unless some mecha- 
nism to concentrate them exists. This might be accom- 
plished by the animal's own swimming reactions. 

G.F. PAPENFUSS: It is known that in temperate seas the 
larger algae are very important in providing protection, 
or a home as it were, for many kinds of animals and thus 
contribute directly to the concentration of life in shallow 
waters. Is this also true in tropical regions where the algae 
are much smaller? 

M.S. DOTY: This is true also in the tropics and parti- 
cularly among the coralline algae which provide many 
niches and serve for protection of small animals. 

H.M. HURKHILL: What species of the mangrove com- 
plex has been introduced to Hawaii for purposes of land 
formation ? 

M.S. DOTY: It is only the Rhizophora spp. 

H.M. DURKIUIL: The primary pioneer mangrove in 
Malaya is considered to be Avicennia spp., but in the man- 
grove areas of Singapore a Dictysta sp. looks as if it takes 
an active initiation in the binding of mud found in the 
intertidal zone and fiats below the mangrove limit. With- 
in the tidal prawn ponds of the mangrove belt, Vaucheria 
appears to be an important mud-binder which may criti- 
cally limit the commercial life of the ponds. 

G.F. PAPLNFUSS: Can it be that the gradual sloping 
of the reef on the windward side of an atoll is owing to the 
scouring action of the waves on that side as compared with 
the conditions on the leeward side? 

F.R. FOSBERG: It should be pointed out that the erosion 
on the slopes of atoll reefs is of two sorts - that by ordinary 
waves and that by typhoon or storm waves. The latter 
may be much more important ; they come from the south, 
predominantly, and produce different effects than ordinary 
waves. The function of the other encrusting organisms is 
to cement the relatively fragile growth-lattice of the reef 
into a rock capable of resisting ordinary wind waves. 





Department oj Geology ', Colorado School of Mines, Golden, Colorado, U.S.A. 


Reefs ranging in age from the Eocene to the 
Recent are known from a number of islands of 
the Western Pacific, and they have been encoun- 
tered in borings made in several of the islands. 
It has been the author's good fortune to have 
spent much of the last ten years studying the fossil 
algae from these reefs. The fossils consist only 
of calcareous algae: crustose corallines, articu- 
lated corallines, Halimcda, and a few Dasycla- 
daceae. The non-calcareous forms have not been 
preserved, so any comparisons between Recent 
and ancient floras will have to be limited to cal- 
careous algae. 

Unfortunately Recent calcareous algae have 
received relatively little study. They are probably 
the least known of any of the algal groups. 
Possibly this is because they are more difficult to 
collect, being commonly firmly attached, and 
because of the necessity of decalcification, section- 
ing, and study with a microscope. 


The Recent Calcareous Algae found in the 
Tropical Pacific appears to form a surprisingly 
uniform widespread flora consisting of a relative- 
ly small number of genera and species. Common- 
ly at a given locality there may be great numbers of 
individuals of a few species. The magnificent 
algal ridge at Bikini has been built largely by two 
species: Porolithon gardineri and Porolithon 
onkodes, while the great Lithothamnion bank of 
Haingsisi near the Southwest point of Timor, 
so vividly described by Weber Van Bosse (77, 
pp. 4-5), is composed largely of rounded masses of 
Lithothamnium eruhescens. Most of the common 
species are surprisingly widespread, extending 
from the East Indies over the tropical Pacific 
and westward across the Indian Ocean. Several 
species extend from the Red Sea to the Marshall 
Islands and even to Hawaii. Dawson records 
some species from both the coasts of Vietnam 
and western Mexico. 

Very few comprehensive listings of the species 
found in a given area are known to the writer. 


However, Table 1 will give an idea of the general 
complexion of the Recent Flora. 


The oldest reefs of the Western Pacific for which 
we have much information are of Eocene age. 
The author has made detailed studies of large 
collections of Eocene material from Saipan (10) 
and the Eniwetok cores ( 11), and is now studying 
a large collection from Guam. He has also stud- 
ied small collections from the East Indies and 
from Ishigaki in the southern Ryukyu Islands. 
Ishijima has described Eocene species from 
Taiwan, and adjoining areas. 

The Eocene flora is another relatively small 
homogeneous flora with a wide distribution. A 
number of species, or very closely related forms, 
have been found from the western Mediterranean 
to the Mariannas and even to the Marshall Is- 
lands. Data showing the general composition 
of the Eocene flora of crustose coralline algae is 
shown on Chart 2. 

The known Miocene flora is considerably 
larger than the Eocene flora, with a greater 
representation of species belonging to the genera 
Lithophyllum and Mesophylluw, fewer species of 
Archaeolithothamnium, and about the same num- 
ber of Lithothamnium. This flora also has a 
number of links with the Western Mediterranean 
and even some with the West Indies. 

The Pleistocene Flora is not well known. 
Much more collecting and study is needed. 
However, it appears to be essentially the same as 
the Recent. The only difference noted so far is 
that in some localities it may contain more 
species than the Recent. This may merely mean 
incomplete knowledge of the Recent. Most of 
the Pleaistocene species are still found growing 
along the present coastlines. The Pleistocene 
flora is characterized by the presence of large 
quantities of Porolithon, Goniolithon, and Amphi- 
roa. In some regions (Palau, Saipan, Bikini, and 
Eniwetok) there is also an abundance of Halimeda 
debris. The Lithothamnium and Lithophyllum 
present usually belong to thin encrusting species. 
Archaeolithothamnium occurs, but it is represented 
by only a few rarely found species. 



Too little is known about the Pliocene Flora 
to merit a discussion here. Charts showing 
details of the composition and distribution of 
these floras will be published in the author's 
forthcoming report on the fossil algae obtained 
from the deep drillings at Eniwetok, Funafuti, 
and Kita-Daito-Jima (11), which will probably 
appear during 1958. 


No reefs older than Eocene arc known in the 
Western Pacific region. Reef building corals and 
rudistids have been dredged from the tops of 
Guyots in the areas around the Marshall Islands 
(1), but no algae were described. Permian reef 
building algae have been found in Timor and 

Table 1. 
Composition, Recent Floras of Calcareous Algae. 

The numbers indicate the number 
of species of each genus. 










Jan ici 




Borne te I la 





Bikini & Northern 
Marshalls (Taylorl950) 


-C x 

Js 8 

**. rH 

C C 

- o 

-3 5 




r 1 


2 o 

0) 7) 

^ s 

5 * 

u B 



S ^ 
r> ^ 

. 8 

bo ;-. 

^ s 


S B 

. ~ ~H 

S S2 
1 1 



to u 

(D (0 

o c 55 


>- u.2 


T^ ^a w 

X w o 
























































Japan. However, it is necessary to go to distant 
continents to learn the character of the algal 
floras of the Mesozoic and Paleozoic Reefs. The 
Mesozoic reef floras are almost unknown. The 
Paleozoic floras have an entirely different com- 

plexion consisting largely of calcareous green 
algae. Calcareous red algae appear to have been 
rare before the Jurassic. During that period they 
were represented mainly by members of the ex- 
tinct family of the Solenoporaceae. The Coralline 

Table 2. 
Geographical Distribution of Eocene Coralline Algae found in the Western Pacific. 

Genus and Species 


A. cf. A. chamorrosum Johnson 

A. dalloni Lemoinc 

A. cf. A. hemchamlri Rao 

A. numniuliticum (Giimbcl) Rothplet/ 

A. oulianovi Pfcnder 

A. a IT. A. saipancnsum Johnson 

A. cf. A. sociahile Lemoinc 

L. cf. lingusticum Airoldi 

L. cf. L. uhraidi Lemoine 

L. crisputhallum Johnson 

L. kumhecrustwu Johnson 

L. cf. nwreti Lemoine 
L. tcipachawn Johnson 

M. robustus Johnson 

M. vaughanii (Howe) Lemoine 


C. prisca Johnson 

Eniwetok | 





























































algae (Family Corallinaceae) are not common 
before the beginning of the Tertiary. Most of the 
genera of today extend back into the Eocene, 
only a few into the Cretaceous. No represen- 


tatives of Recent genera have yet been found in 
rocks older than middle Jurassic. The known 
geologic range of the common genera of the 
Corallinaceae is shown in Table 3. 

Table 3. 
Known Geologic Range of Common Recent Genera of Coralline Algae. 













(1) Barton, E.S., 1901, The Genus Halimeda: 

Siboga Expedition Monograph, 60: 1-32, 
pbs. 1-4, Leyden, Holland, Brill. 

(2) Dawson, E. Yale, 1954, Marine Plants in 

the Vicinity of Nha Trang, Viet Nam, 
Pacific Science, 8:373-469, 63 figs. 

(3) , 1956, Some Marine Algae of 

the Southern Marshall Islands, Pacific 
Science, 10:25-66, 66 figs. 

(4) , 1957, An Annotated List of Ma- 


































































































rine Algae from Eniwetok Atoll, Pacific- 
Science ; 11:92-132, 31 figs. 

(5) Foslie, M., 1900, Calcareous Algae from 

Funafuti, Del. Kgl. Norske Via 1 . Selsk., 
Skrifter, 1900, 1:3-12. 

(6) Foslie, 'M.H. and Printz, H., 1929, Contri- 

butions to a Monograph of the Litho- 
thamnia, Det. Kongl. Norske Vid. Selsk, 
Museet, Monograph, 60 pp. 75 pis. 

(7) Hamilton, E.L., 1956, Sunken Islands of the 

Mid-Pacific Mountains, Geol. Soc. 
America Mem., 64, 97 pp., 12 figs., 13 pis. 



(8) Ishijima, W., 1954, Cenozoic Algae from 

the Western Pacific, Tokyo ^ Yuhodo 
Company, 87 pp., 69 pis. 

(9) Johnson, J. Harlan, 1954, Fossil Calcareous 

Algae from Bikini Atoll, U.S. Ceol. 
Survev Prof. Paper 260-M, P. 537-545, 
pis. 188-197. 

(10) _, 1957, Fossil Calcareous Algae 
from Saipan, U.S. Geol. Survey Prof. 
Paper 280, Part E, pi. 25 (in press). 

(11) _., 1958, Fossil Calcareous Algae 
from the Eniwetok, Funafuti, and Kita- 
Daito-Jima Drill Holes, U.S. Ceo*. 
Survey Prof. Paper 260-W (in press). 

(12) Johnson, J. Harlan and Ferris, B.J., 1949, 

Tertiary Coralline Algae from the 
Dutch East Indies, Jour. Paleontology, 
23, (2): 193-198, pis. 37-39, (March). 

(13) Lemoine, Mme. P., 1939, Lcs Algucs Cal- 

caires Fossiles de L'Algerie, Mat. 

Carte Geol. de L'Algerie Series, 9, 
128 pp., 3 pis., 80 figs. 

(14) Lignac-Grutterink, L.H., 1943, Some Ter- 

tiary Corallinaceae of the Malaysian 
Archipelago: Verk. Geologisch-Mijn- 
bouw Kundig Genootschap veer Neder- 
land en Kolonien, Geol. Ser., Deel, 13: 
283-297, 2 pis., (Dec.). 

(15) Sripada, Rao K., 1943, Fossil Algae from 

Assam: 1 The Corallinaceae, Proc. Nat. 
UAcad. Sciences, India, 13, part 5:265- 

(16) Taylor, W.R., 1950, Plants of Bikini and 

other Northern Marshall Islands: Uni- 
versity Michigan Press, 227 pp., 79 pis. 

(17) Weber van Bosse, A., and Foslie, M.H., 

1904, The Corallinaceae of the Siboga 
Expedition: Siboga, Exp. Monograph 
LXI, 1 10 pp., 16 pis., Leyden, Holland. 


M.S. DOTY: Can Halimcda be recognized in fossil depo- 

j.i i. JOHNSON: Halimcda can be recognized to genus 
quite readily, but it is not as easy to compare with species 
because parts used by the taxonomist dealing with modern 
species may not be preserve in the fossil record. They are 
quite abundant and contribute a great deal to the shallow 
water deposits back as far as the Cictaceous. 

j.v. SANTOS: Which plays the greater role in the forma- 
tion of an atoll, the coelentcrate corals or the coralline 

J.H. JOHNSON: It may vary to some extent, but in 
general the formation of the atoll is made up basically of 
coral and then the coralline algae come in, with diatoms 
and foraminifera finally filling in the spaces. 





Pacific Vegetation Project, Care of National Research Council, Washington* D.C., U.S.A. 

A coral atoll may be described, in the briefest 
terms, as a cap of limestone of organic origin on 
a mountain on the floor of the ocean, rising to or 
only slightly above sea level. Some lie on shallow 
banks or continental shelves. The cap is a ring- 
like ridge or reef surrounding a body of water 
termed a lagoon. Some parts of this reef may 
emerge above high tide level as islets. These may 
either be remnants of former higher reef levels or 
detrital accumulations. Much or all of the reef 
surface below mean low-tide level and down to 
depths where sun-light penetration is very atten- 
uated is composed of communities of living 
plants and animals. In bulk, at least, these are 
mostly organisms that secrete limy skeletons. 
Accumulations of these skeletons make up, 
almost exclusively, the reefs and upper parts of the 
mountain down to the volcanic (or other) base- 
ment rock on which the reefs originally started 
to grow. The depth of this limestone is known 
for only a few atolls and may vary from at least 
1 ,400 meters to, probably, very much less. 

The concept of the ecosystem, first proposed by 
Tansley (9), is that of an interacting system com- 
posed of an environment and all of the organisms 
involved with it. It is normally an open system 
because there is a continuous, though variable, 
influx and loss of energy and material. Such a 
system is, of course, an abstraction constructed 
to facilitate understanding of the complex pro- 
cesses involved in a segment or class of segments 
of the biosphere. As such, its extent is limited 
only by selection and definition of the segment 
or segments under study. Thus it may be varied, 
in different examples, from the smallest observa- 
ble unit of environment in which organisms live 
to the entire world's biosphere as a whole with its 
total environment (4). As the ecosystem is 
limited only by the extent of the effective environ- 
ment, the maximum could be, theoretically, the 
entire universe. Practically, however, the defini- 
tion will not ordinarily extend to the ultimate 
sources of energy, or even of material. It will be 
restricted to such extent as will best facilitate 
observation and understanding of the portion 
of nature under immediate study. This concept, 
of obvious and increasing utility but not too easy 
to handle, and never, apparently, used by its 
creator, has been, in recent years, adopted by a 

number of ecologists (e.g., 6, /, 5, 4, 2, 3). No 
two have defined or formulated their ecosystems 
in exactly similar terms, nor is there any critical 
need, at this stage, to do so. 

In this paper, the coral atoll ecosystem will be 
described in terms of processes involving transfer 
or transformation of energy and material, with 
only incidental reference to the actual organisms 
involved in the system or to the physical struc- 
tures found in the environment. It is recognized 
that in a complete account of such a system these 
aspects, also, would be described fully. For 
present purposes it may suffice to say that the 
biotic component of the system is composed of 
phytoplankton and zooplankton; free-living but 
bottom-dwelling animals and other hctcrotrophic 
organisms of many sorts; an enormous aggrega- 
tion of sessile or fixed organisms representing 
most classes of algae, a few seed plants, and prac- 
tically all phyla of invertebrate animals; and a 
diverse assortment of land animals and plants. 
Many of the marine organisms secrete skeletons 
of CaCO 3 which arc added to the material of the 
substratum. This process forms a lattice-work 
of limestone in which free skeletons or loose 
fragments lodge. By several processes these 
may become bonded in such a way as to form a 
rather hard and resistant rock. This is built up in 
the form of a ridge or reef usually enclosing a 
shallow body of water or lagoon and rising 
variously to somewhat below, near, or just above 
high-tide level. This reef is ordinarily rather 
flat-topped, of varying width, with irregularities 
or islets extending above high-tide level. Waves 
commonly break on the outer margin and water 
flows from the sea to the lagoon and back to the 
sea over the flat or gently sloping reef surface or 
through gaps in it. The flow may be in and out 
with the tides, or in over the windward and out 
over the leeward sides. The islets are commonly 
composed in part or wholly of loose porous 
limestone debris and are mostly covered by 
vegetation that includes representatives of all 
major groups of land plants. The land faunas 
are made up of a large number of species of 
insects and other arthropods, worms, land 
molluscs, a few reptiles, some birds, and a few 
mammals, including rats and man. Larger islets 
may contain within their porous structures a body 



of fresh ground water floating on the underlying 
salt water and retarded by friction from free 
diffusion with it. 

These atoll structures are found in most regions 
of the tropical and, more rarely, subtropical seas. 
Temperatures range generally between 75 and 
85F or in full sun on land, higher, decreasing 
of course with increasing depth in the sea. 
General climates range from relatively dry, 
perhaps 600 mm precipitation, to wet, 5,000 mm 
or more. The atolls are in trade wind, doldrum, 
and monsoon belts. Insolation is generally high, 
cloudiness low to moderate. 

Most atolls are inhabited by human beings, 
some by relatively large populations. These 
exert a generally appreciable, often profound, 
influence on the functioning of the ecosystem. 
Of specific importance in this connection are the 
economic activities of planting coconuts, harvest- 
ing, drying, and exporting coconut meat, and 
importing in exchange various foods and other 

With this description of the general physical 
and biological situation, we may proceed to 
describe in more formal terms, the abstraction 
called the coral atoll ecosystem. This may be 
outlined as follows in 12 sections, lettered A to L. 

A. Media. 

B. Nourishment or inflow. 

C. Production. 

D. Transformation. 

E. Decomposition. 

F. Excretion. 

G. Accumulation. 
H. Turnover. 

I. Miscellaneous other elTects and processes. 
J. Losses. 
K. Balance. 
L. Trends. 

A. Media: The media in which the system 
exists are two a layer of sea water and a super- 
imposed layer of air. These, by the nature of 
the earth-system itself, are constantly changed by 
sea and air circulation. Through them or by 
means of them, all exchange, gain and loss, of 
matter and energy takes place. These two media 
are the most universal and pervasive components 
of the ecosystem and at the same time its environ- 
ment, influencing in some measure everything 
in the system. 

B. Nourishment or inflow: As the atoll is an 
open system, there is a continuous addition of 
matter and energy in many forms. Fundamental, 
of course, is the daily increment of solar energy 


without which the system, in anything like its 
present form, could not exist. Its functioning is 
in almost every respect dependent on either photo- 
synthetic or thermodynamic processes, which are 
dependent on constant addition of energy from 
the sun. The nourishment of organisms and the 
circulation of both air and water are important 

The energy of wind, mostly indirectly a form of 
solar energy, also exerts its force in various ways 
in the system. Most of this energy is received 
elsewhere and transported to the atoll. 

The gravitational energy of both sun and moon 
contributes importantly to sea-water circulation 
in the form of tides. The movement of ground 
water in atoll islets also is influenced by tidal 

Essential components of the media, such as 
O 2 , N 2 , H 2 O, CO 2 , as well as dissolved salts and 
suspended organic matter, and even living pro- 
pagules and disseminules of organisms are con- 
tinually renewed or carried into the system by air 
and ocean currents. Relative concentrations of 
the various components of the media are main- 
tained at a rather constant level in this manner. 
The replenishment of the ground-water bodies in 
islets is dependent on incoming fresh water, 
mostly evaporated elsewhere and deposited as 
rain on the islets from wind-borne clouds. 
Surface currents, upwellings, tradewinds, mon- 
soons, and cyclonic storms are important aspects 
of the circulation patterns involved. The intro- 
duction into the system of phosphorus, on which 
organic activity is completely dependent, is 
believed to be controlled to a considerable extent 
by upwellings of deep-sea water. One important 
route of transport of phosphorus from areas of 
upwelling to the atolls is by means of fish-eating 
birds and their young which deposit phosphates 
in their excreta within the area of the system. 
Essential mineral elements, nitrogen, and organic 
carbon are also brought in by the birds at the same 
time. Organic matter, in the form of drifting 
plankton, driftwood, and dead organisms, is 
brought to the atolls by currents. Currents also 
bring small amounts of mineral elements in the 
form of pumice as well as in solution. Volcanic 
ash arrives by way of winds, especially high alti- 
tude winds. 

Finally, with changing patterns of human 
activity on atolls, increasing amounts of imported 
foods and other materials as well as alien organ- 
isms are introduced into the system. These 
introductions are effecting various rather pro- 
found changes in the equilibria and altering 
greatly the physical appearance of the atolls. 



C. Production: The elaboration by plants of 
basic organic materials from elements and simple 
inorganic compounds is termed production, in 
an ecological sense. Such elaboration provides 
the fuel and building materials for all other life 
processes. The effective capacity of a system for 
production is called its productivity. (1) The 
most obvious productive process is photo- 
synthesis, by which carbohydrates arc elaborated. 
Algae and green plants utilize CO 2 , H 2 O, and 
energy from sunlight for this purpose. Such 
algae occur as components of plankton, within 
the cells of corals and other coelenterates, fixed 
on the reef surfaces, terrestrially on soil and rocks, 
and epiphytically on tree trunks. Mosses are 
found in many terrestrial situations on the islets, 
as well as on tree trunks, especially where they 
are shaded. Ferns and psilopsids are common 
growing on land and epiphytic on trees. Seed 
plants grow principally on land, but some are 
epiphytic and a few are marine, growing in shal- 
low lagoon situations on sandy bottoms. All of 
these groups, together, account for the origin of 
most of the carbohydrates used in the system. 
(2) The other essential type of production is the 
fixation of atmospheric nitrogen its oxidation 
and elaboration into simple compounds. It is, 
well known that this fixation is accomplished by 
bacteria in the soil and in nodules on the roots 
of certain leguminous plants. Less well known, 
but possibly more important in the atoll system, 
is fixation of nitrogen by blue-green algae. This 
occurs on the soil surface and possibly in fresh and 
salt water. Much of the nitrogen available to 
atoll organisms is probably fixed within the system, 
but important quantities enter the system by way 
of birds, rain, and ocean currents. 

D. Transformation: The alteration oj primary 
and fabrication of secondary organic compounds: 
This function may be viewed as a succession of 
processes, mostly involving a break-down of 
organic compounds and their re-elaboration into 
more complex ones. Some of these are of an 
enormous order of complexity (e.g., nucleo- 

(1) Autotrophic plants, in the nourishment of 
their own protoplasm and elaboration of stored 
material, cellulose, lignin, and other materials, 
carry out the first major step in a series of 
turnovers of the products of photosynthesis. Of 
course, additional inorganic materials are incorpo- 
rated by this process, and many of the elaborated 
compounds are infinitely more complex than the 
original carbohydrates produced by photosyn- 

(2) Heterotrophic (parasitic and saprophytic) 
plants carry this process a step farther in utilizing 
already elaborated complex substances, as well as 
simpler materials derived from their hosts and 
organic substrata. Here may be mentioned the 
utilization of dissolved organic matter in the 
media by facultatively heterotrophic planktonic 
algae as discussed by Saunders (8). 

(3) Animals, feeding on plants in various ways, 
convert plant organic matter into animal organic 
matter. The principal classes of processes by 
which this is accomplished are as follows: 

a. Eating of phytoplankton by zooplankton. 

b. Utilization of material produced by zooxan- 
thellae, by their coelenterate hosts. 

c. Reef grazing and boring. 

d. Eating of land plants by animals. 

e. Eating of dead plant parts by animals. 

f. Parasitism of plants by animals. 

(4) Secondary conversion of animal matter to 
animal matter is accomplished as a result of 
three well-known classes of processes, namely: 

(a) Predation 

(b) Parasitism 

(c) Scavenging. 

These are carried on in a great number of different 
ways by a large number of animals. Reef grazing 
and boring are important here, too. 

(5) Reconversion of animal matter to plant 
matter is not as conspicuous a process, but is 
important nevertheless. There seem to be no 
insectivorous plants on atolls, so this reconver- 
sion is principally accomplished by bacteria and 
fungi living mostly on dead, and occasionally on 
living organic matter. It is an interesting question 
whether zooxanthellae utilize in any way the 
materials of their hosts' tissues. 

E. Decomposition (usually but unfortunately 
termed "reduction"): The destruction of the 
elaborated organic compounds and reconversion 
back to simple inorganic compounds and relative- 
ly inert organic residues: Two main categories 
of processes are involved here. (1) Physiological 
oxidation (inappropriately termed respiration by 
many plant physiologists), which is the oxida- 
tion of carbohydrate materials within living cells 
releasing the energy required for other life pro- 
cesses. This process goes on constantly in all living 
things. (2) Non-biological oxidation, both by 
burning and by the slow oxidation of dead materials 
that normally takes place on exposure to atmos- 
pheric oxygen, aided or not by hydrolytic and 
catalytic action. 



The principal products of both sorts of processes 
are CO 2 and H 2 O, with, of course, inorganic 
and inert organic residues. Chemical energy is 
released and converted into other forms. 

F. Excretion (within the system): The release 
of waste products and residues by organisms into 
the media: 

(1) In water, CO 2 and O 2 are released, as well 
as excreta and soluble metabolic wastes. Calcare- 
ous and siliceous skeletons and oily material 
remain after disintegration of organisms. 

(2) On land, likewise CO 2 , O 2 , and metabolic 
wastes arc released in solution in air or water. 
Guano and other excreta, as well as deciduous, 
caducous, or severed plant parts are deposited 
on the surface of the ground to decompose. 

G. Accumulation- Storage of materials in 
unchanging or very slowly changing form, i.e., 
temporary withdrawal of material (and energy) 
from free circulation in the system. 

(1) In bulk the limestone from calcareous 
skeletons of plants and animals represents the 
greatest and most important accumulation, the 
principal component of the atoll itself. 

(2) Phosphatic residues, mainly calcium phos- 
phates, are present as phosphate rock, compo- 
nents of soils, guano, and at least in some atolls 
(e.g., Washington Island) as a phosphatic mud 
or sludge on the lagoon bottom. 

(3) Humus, both as raw humus in Pisonia 
forests, and as more stable humic residues in Aj 
horizons of soils, plays an important part in the 
functioning of the system. The acid raw humus 
contributes to the formation of phosphate rock, 
and the soil humus helps to maintain the soil 
in condition to support growth or larger plants 
and micro-organisms. Humus, though relatively 
inert, is continually undergoing a slow oxidation. 

(4) Slight accumulations of charcoal, metal 
oxides, silica, and silicates occur where human 
activity is significant. Silica from sponge, 
radiolarian, and diatom skeletons also occurs in 
minute amounts, as well as small quantities of 
silicates from floating pumice. 

(5) Finally, fresh water, in the ground-water 
lens, as well as in the several states of soil water, 
may be regarded as a temporary accumulation. 

The organic matter and other substances in 
living organisms represent a large total quantity 
but, as they are in a constant state of turnover, 
should probably not be classed as an accumula- 

These accumulations, along with the materials 
in solution in the media, may be regarded as the 


reservoirs of materials on which the other com- 
ponents may draw for nourishment. 

H. Turnover of materials and energy: Cate- 
gories D, E, and F, above are to be classed as 
turnover. In addition several more processes 
may be so regarded. 

1. Re-use of CO 2 released by oxidation. 

2. Re-use of O 2 released by photosynthesis. 

3. Re-use of H 2 O released by metabolic and 
external chemical processes. 

4. Re-use of fixed nitrogen, both from meta- 
bolic wastes and from primary biological oxida- 

5. Re-use of mineral nutrients released by 
excretion and breakdown of organic materials. 

6. Withdrawal from and return of various 
materials to media. 

7. Withdrawal from and return to accumula- 

I. Miscellaneous other effects and processes 
taking place within the system. 

1. Inhibition by salt (NaCl). The organic activ- 
ity in terrestrial situations seems subject to a 
considerable inhibition by the salinity of the sea- 
water medium. This inhibition results from 
difference in osmotic pressure, the chemical 
effects of absorption of excess sodium and chlo- 
rine ions and consequent inhibition of absorp- 
tion of others. The number of land organisms 
completely adapted to the normal salt concen- 
tration of the sea is limited. Hence establishment 
of immigrant organisms is severely limited, and 
many of those that become established function 
at below their optimum levels. Salt water enters 
the land environment as wind-borne spray, as 
storm waves, and by diffusion through the 
ground. The conspicuous nature of the limiting 
effects of salinity may be a reflection either of the 
small extent of the land habitat and consequently 
great exposure to salt or of its probable geolog- 
ically recent origin that has allowed little time 
as yet for evolution of a special atoll biota. 

2. Effects of sea-air interface: Category 1 is 
really only one of the consequences of the fact 
that the land portion of this ecosystem is a thin 
lens inserted in the general sea-air interface. 
The distribution of many organisms, marine as 
well as land, is influenced by the character of this 
interface. Aeration, principal release of energy 
from insolation, frequently an abrupt break in 
temperature gradient, local high salt concentra- 
tions resulting from evaporation, solution and 
other forms of erosion of limestone, and the 



shaping of the contours of vegetation and con- 
trol of its composition are all consequences of 
the nature of this interface. Many more could 
be enumerated. 

3. Shelter effects. One of the reasons for the 
diversity of animal life in such an apparently 
simple environmental complex may be the 
variety of habitats resulting from the surface 
irregularity of the several substrata. The vegeta- 
tion, the deeply pitted rock, the porous soil, and 
the intricate nature of the reef lattice provide 
shelter of various types for a large number of 
species of animals (and plants, too) that have 
widely differing requirements. 

4. Burrowing and turning over of soil by crabs 
is an important factor in the process of incor- 
porating organic matter into the soil. Crab 
burrows are very common on many atolls, and 
fresh mineral soil is often piled or scattered 
around their entrances. The mechanical tilling of 
the soil in this manner has been compared to that 
accomplished by earthworms in other habitats. 
It doubtless is a process of great importance, 
though no careful assessment of its extent or 
effects has been made. 

J. Losses (or excretion from the system): 
a. Of the principal substances lost from the 
system the first three listed below are present in 
such constant proportions in the media outside 
the ecosystem that the losses may be considered 
as balanced almost exactly by inflow. The others 
are fluctuating quantities, and there is no exact 
relation between inflow and loss. 

1. CO 2 , carried away by winds and currents. 

2. O 2 , carried away by winds and currents. 

3. N 2 , carried away by winds and currents. 

4. Fresh water dispersed into media and carried 
away by winds and currents. 

5. Nitrates and organic N, carried away by 

6. Phosphates, carried away by currents in 
solution and suspension. 

7. Other dissolved mineral substances, carried 
away by currents. 

8. CaCO 3 , carried away by currents in solu- 
tion and suspension. 

9. Plankton, carried away by currents. 

10. Dead animals and plants and detached living 
fixed organisms, carried away by currents and 

11. Birds which migrate. 

12. Export of copra. 

13. Export of pearl shell, etc. 
b. Energy losses: 

1. Light, by reflection. 

2. Heat, by radiation and convection and 
carried by winds and currents. 

3. Chemical energy lost with elaborated mate- 

K. Balance: The resultant of all of the factors 
at work on the segment of the universe (or of 
nature) occupied by the atoll ecosystem is the 
atoll itself. It may be thought of as a system 
in a state of dynamic equilibrium with a positive 
offset represented by the physical mass of the 
atoll with its associated biota and its total of 
organic and mineral matter over and above that 
of the normal media air and sea water that 
otherwise would occupy the space. All the 
characteristics described serve to set the system 
off from the surrounding undifferentiated media. 

L. Trends: With such complexity, it is hard to 
estimate trends, though it may be easy to discern 
them. Over very long periods, the trend is obvi- 
ously toward greater accumulation of material 
and probably toward increasing complexity. 
This trend usually seems directly related to slow 
subsidence of substratum on which the atoll is 
built, and may be expected to continue. On a 
shorter time scale, it is possible to suggest that 
during periods of general or eustatic rise in sea 
level mass will increase, by addition of calcareous 
material in layers. Biotic complexity may at the 
same time decrease somewhat with tendency 
toward loss of land habitats. With fall in sea level, 
the trends may be the opposite loss of mass by 
erosion and gain in biota with appearance of land 
habitats, increased activity of sea birds, and 
especially the results of occupation by man. 
Presumably for about the last 3,500 years the 
latter trend has been generally maintained. 
Whether or not the last few decades have wit- 
nessed a change in this trend is uncertain. 

It seems clear that these major trends are 
controlled by factors external to the system. The 
ultimate control of sea level is as yet by no means 
clear. The variation in CO 2 content of the air has 
been suggested (7) as a factor that determines, 
or at least influences, world temperatures, eva- 
poration of sea water, accumulation of ice, and 
consequent effects on sea level. It has been sug- 
gested that the recent apparent reversal of the fall 
of sea level may be due to the vastly accelerated 
industrial activity which pours great quantities of 
CO 2 into the atmosphere. If this is a valid 
assumption it seems reasonable to think that the 



present rise will continue, probably at an increas- 
ing rate. Thus a prediction might be made that 
the presently observed loss of land above water 
by erosion may be accelerated by a rise in sea 
level and consequent submergence of much or all 
of the land area of atolls. Such predictions, 
however, rest on very insecure bases at present. 

On a still shorter time scale are the effects 
produced by the occupancy of the atolls by man, 
and especially modern man. These effects tend 
to be drastic as far as the land portions of the 
ecosystem are concerned, but trends arc as yet 
hard to isolate. Certainly the replacement of the 
native vegetation by coconut plantations and the 
rise of the export of copra are notable and 
probably involve a complex of related or depen- 
dent effects. This change will probably continue 
but certainly at a decelerated rate, as land for 
expansion of plantations is becoming scarce. 
Augmentation of the land biota will probably 
continue as man's effect on the land environment 
continues. Pollution of lagoons will undoubt- 
edly increase, with resulting encouragement to 
some organisms and ill effects on others. Fishing 
activities have tended to decrease with contact 
with civilization, but this trend may probably be 
reversed and with use of such effective methods as 
dynamiting and poisoning the marine biotas may 
undergo considerable change. There has as yet 
been little attempt to measure the results of such 
factors so that here, as in other aspects of the 
system, estimation of trends is highly speculative. 
If such prediction is of interest, attention should 
be directed toward critical study of the details 
of the working of the system outlined above, to 
clarify it and fill in the parts that are at present 
inferential. It seems possible that if a firm under- 
standing of this ecosystem is achieved it may be 
used as a model in terms of which to study other 


The general physical and "physiological" 
framework of the coral atoll ecosystem has been 
outlined in terms of the media in which the system 
exists, nine categories of processes taking place 
within the system, the balance or dynamic equi- 

librium in the resultant structure brought about 
by these processes, and suggested trends in the 
state of this equilibrium. This highly generalized 
picture rests on a vast accumulation of facts and 
upon inferences drawn from them and from 
pertinent facts derived from study of related or 
analogous situations in other systems. It is hoped 
that this description may serve, until a better 
conception is devised, as a framework around 
which an understanding of this segment of nature 
may be built and as a guide for future research 
designed to clarify our knowledge and apprecia- 
tion of coral atolls. 


(1) Billings, W.D., and Bliss, L.C., 1955, An 

Alpine Snowbank Ecosystem in the Medi- 
cine Bow Mountains of Wyoming, 
Bull. Ecol Soc. Amer., 36: 76 (abstract 

(2) Cain, S.A., 1956, The Expansion of the 

Human Ecosystem, 1-10 (duplicated, 
distributed at Symposium: Values in 
human ecology, New York, Dec. 28, 

(3) Dansereau, P., 1957, Biogeography, An Eco- 

logical Approach, 1-394, New York. 

(4) Evans, F.C., 1956, Ecosystem as the Basic 

Unit in Ecology, Science, 123 (3208): 

(5) Costing, H.J., 1956, The Study of Plant 

Communities, 1-440, San Francisco. 

(6) Pitelka, F.A., 1955, High Arctic Tundras as 

an Ecosystem, Bull. Ecol. Soc. Amer., 
36: 97 (title only, paper read at East 
Lansing meeting of Ecological Society 
of America, Sept. 1955). 

(7) Plass, G.N., 1956, Carbon dioxide and the 

Climate, Amer. Scientist, 44: 302-316. 

(8) Saunders, G.W., 1957, Interrelations of Dis- 

solved Organic Matter and Phytoplank- 
ton, Bot. Rev., 23: 389-409. 

(9) Tansley, A.G., 1935, The Use and Abuse of 

Vegetational Concepts and Terms, Eco- 
logy, 16: 284-307. 


F.E. EGLfcR*. The approach used by the author and its 
potential as an approach to land utilization is most valu- 
able. The concept of the ecosystem has been slow to gain 
recognition, possibly because it is based on the investiga- 
tions of many fields of knowledge, seldom as the result of 


one individual's effort. The plant ecologist often comes 
from the field of taxonomy and may not be fully equipped 
to handle phytosociological problems. The integration of 
scientific knowledge by an ecologist requires an unusual 
breadth of understanding. 


M.S. DOTY : It is necessary to distinguish between poten- regarding when it stops being an atoll and when it becomes 

tial and net productivity. a lagoon. With reference to the energy relationships, the 

F.R. FOSBERG: Although potential productivity is some- constant blowing of wind corning from outside may contri- 

thing to think about and discuss, the actual productivity bute to the system, but the chief source of energy is from the 

is the only tangible aspect of it that could be evaluated. sun. The wind may build up sand and accumulate energy 

G.F. PAPENFUSS: Are there atolls that lack a lagoon? in the system, but it may also do the opposite and release 

F.R. FOSBERG i There is some question of interpretation potential energy from the system. 





School of Fisheries, University of Washington, Seattle, Washington, U.S.A. 

In the U.S. Pacific Northwest, one is accus- 
tomed to seeing large fronds of such algae as 
Macrocystis and Ncrcocystis floating around in 
the cold water of Puget Sound. In the Marshall 
Islands of the warm Central Pacific Ocean, on 
the other hand, where the species of algae dis- 
cussed in this paper were collected, the absence 
of these or any of the large forms of algae is 
very apparent. There only colonies or individual 
specimens of tiny filamentous algae or branched 
species rarely exceeding a few inches in length 
were found. The striking size difference in the 
marine flora of the two regions brings up the 
question as to the nature of the conditions caus- 
ing this difference, and, at the same time, poses 
the question as to reasons for the differences in 
distribution of the algae in the atoll itself. This 
paper will be confined to the latter problem. 

A coral atoll is a relatively small outcropping 
in the middle of an ocean and contains a diversi- 
fied environment in which the growth of various 
organisms is possible. The open ocean surround- 
ing an atoll is a source of the minerals used in 
the production of organic matter and thus acts 
as a nutrient solution for the activities of the 
various photosynthetic organisms present. The 
principal members of the atoll community engag- 
ing in photosynthetic activity, of course, are the 
benthic and the planktonic algae. The distribution 
of planktonic algae of the atoll lagoon is primarily 
dependent on the local currents, and con- 
sequently, the problem of this paper is to deter- 
mine the differences in the environment which 
might account for the distribution of the benthic 
algae about the atoll. 

Recent studies by Johnson (5) and others have 
confirmed earlier suggestions that the calcareous 
algae play an extremely vital part in the building 
of coral atolls. Transects across the reef at 
Bikini Atoll indicate that 40-90 % of the reef is 
comprised of algal deposits, consisting primarily 
of the remains of species of the green coralline 
alga Halimcda and of the red coralline algae such 
as PorolithoH, Cioniolithon, Mesophyllutn, and 
Lithothamnion. Other organisms, such as corals 
and foraminifera, have contributed to the reef 
building process, but to a lesser extent. These 
deposits extend to a depth of over 2,500 feet and 

t Presented in abstract by M.S. Doty. 

date back to the Miocene epoch. They serve as 
pointers to the equilibrium or relatively stable 
conditions that exist between the atoll inhabitants 
and their environment. This theory has also 
been promulgated by Odum and Odum (8) who 
stated that the productivity of a reef just balances 
the respiration, suggesting that the reef commun- 
ity is a true ecological climax or open steady state 

The conditions that have existed, apparently 
for millions of years, making possible this reef 
building process, revolve around the activity of 
these algae and their ability to flourish in this 
type of environment. The warm temperatures 
and the abundant radiant energy bathing the 
exposed reefs combined with the rich supply of 
oxygen and carbon dioxide form ideal conditions 
for photosynthesis. The ability of these algae 
to secrete calcium carbonate as well as other 
minerals caused these particular species to sur- 
vive, and their cementing action caused the 
consolidation of algal and other debris in the 
formation of the atoll mass. 

The calcareous algae mentioned above are 
found primarily in the regions of pounding surf, 
where turbulence and oxygen rich waters are at 
a maximum. Some of the non-encrusting cal- 
careous algae, such as Codium and Udotea, 
however, are found in the more protected habi- 
tats of the tide pools and along the western 
leeward reefs where the surf action is less severe. 
These species apparently cannot survive the 
rigorous action of the waves impinging upon the 
northern reefs, but they do require similar condi- 
tions of light, temperature, and mineral compo- 
sition optimum for the growth of the more hardy 

The calcareous algae, however, are not the 
only algae contributing to the mass of the reef. 
Odum and Odum (8) have shown that the fila- 
mentous green algae interwoven in the skeletal 
material of the coral reef make up a large por- 
tion of the mass and consequently increase the 
overall productivity of the reef. 

Of the conditions influencing algal growth in 
an atoll, temperature is probably the least im- 
portant. Studies that showed insignificant 



temperature differences in the shallow waters 
on the different sides of the coral reef were made 
at Rongelap Atoll by Sargent and Austin (9). 
The temperatures across the three hundred- 
meter reef varied from 28.3C on the seaward 
side to a maximum of 29.4'C on the lagoon side, 
not sufficient to account for the differences in 
productivity per unit area on the reef and in the 
adjacent waters. Munk, Ewing, and Revelle (7) 
found very slight temperature differences with 
depth in the lagoon at Bikini Atoll. At three 
feet below the surface, the temperature varied in 
24 hours between 27.2 l 'C and 27.T J C and at 
165 feet between 27.05 and 27.08C, indicating 
a thorough mixing of the top and bottom layers 
of water. This idea was partially supported by 
radiological determinations made after the Baker 
atom test in 1946, by these investigators as well 
as by W.L. Ford (3). The temperatures in the 
atoll environment, then, vary only slightly and 
probably not enough to account for distribution- 
al differences of the organisms present. The most 
significant part that temperature plays undoubt- 
edly occurs during low tides when the tide pools 
become very warm and the beach rock areas are 
exposed to the direct sunlight for hours at a 
time. Here the only algae that survive are the 
blue-green algae such as Anacystis and Schizo- 
thrix, which possess simple structures and gela- 
tinous sheaths. 

Of the factors limiting the growth of photosyn- 
thetic organisms, light is often the controlling 
one. In the murky waters of many northern 
seashores, the stratification of algal species is 
well defined. This zonation can be explained 
at least in part by the amount and quality of the 
incident light and its effects upon the photosyn- 
thetic activity of the various algae. For exam- 
ple, the Chlorophyta are found only in shallow 
water, as the red light from which they obtain 
energy for photosynthesis cannot penetrate more 
than a few feet. Some of the Rhodophyta, on the 
other hand, may be found growing at depths up 
to 300 feet because they can utilize the blue wave 
lengths which penetrate to this depth. The Phaeo- 
phyta use both the red and the deeper penetrating 
orange-yellow light and thus frequent the zone 
extending from the low tide line to a depth 
of approximately 50 feet. Whether such an 
explanation can be used to account for the dis- 
tribution of the algae in an atoll environment is 
open to question. Wells (11) reported that 
"light or radiant energy is the principal factor 
controlling the depth of growth of mutually 
interdependent reef corals and symbiotic zooxan- 
thellae... temperature is the principal factor con- 

trolling their geographic distribution." On the 
other "hand, Sargent and Austin (9) stated in 
Rongelap lagoon, even though the light intensity 
at 15 meters was only 20 " of that just below 
the surface of the water, it was still adequate for 
maximum photosynthesis in free-living plants. In 
the unusually transparent water of Eniwetok 
lagoon at depths up to 45 meters, the present 
author and others of this laboratory using self- 
contained breathing apparatus saw luxuriant 
"fields" of green algae, especially Halimeda 
nionilc, H. gigcis, and Caulerpa raccmosa, attached 
to the white bottom sand. Gilmartin (4) has also 
observed these and other green algae at depths 
of 60 meters. Others have also noted the pre- 
ponderance of green algae at depths usually 
restricted to the deeper living red algae (10). 
This can signify only that sufficient light does 
penetrate the clear waters of these lagoons to 
permit maximum growth of the "greens" and 
possibly to discourage growth of the "reds." It 
was also noted that some of the same species 
inhabiting the deeper, relatively calm water were 
also growing vigorously in shallow turbulent 
water on the edge of the seaward reefs. Good 
examples of this type of algae are the green 
Halimeda opimtia and the red Asparagopsis taxi- 
fonnis. In these instances at least, light does 
not appear to be the controlling factor. 

Another factor important in the growth of 
algae is the mineral content of the medium. 
Few quantitative studies have been made of the 
mineral content of the waters in an atoll environ- 
ment. Sargent and Austin determined the 
amount of dissolved phosphorous in the water 
at several localities on one of the reefs at Rongelap 
Atoll and found that only small differences existed, 
in fact, the values were so low that the measure- 
ments were felt to be uncertain. The total dissolved 
phosphorous over the reef at a depth of about 
forty centimeters varied from 0.31 to 0.66 mg 
atoms/liter. Measurements of Mg++, Ca++, Cr, 
and total hardness made by Cloud (1) at Onotoa 
Atoll in the Gilbert Islands showed that even 
though there were diurnal variations in the 
values for some of these factors, there were no 
significant differences in the levels in the shallow 
water at the four localities studied. Radiological 
studies to determine the relative uptake of trace 
minerals by marine organisms have been con- 
ducted by this laboratory. These studies have 
shown that minute amounts of manganese, 
zirconium, zinc, cobalt, cesium, and iron can 
be detected in amounts too small to be detected 
by ordinary chemical techniques (6). The con- 
centration of these minerals by sedentary marine 



organisms over a long period of time could be 
useful in the measurement of the different minerals 
present and could also be useful in evaluating 
their distribution about the atoll. Until further 
information of a similar nature encompassing 
several ecological situations is obtained, no finite 
conclusions can be drawn regarding the effects 
of varying mineral composition on algal growth 
in the environment of an atoll. 

A factor which must be considered in the 
distribution of an alga is its habit or life form. 
Several classifications of the life forms of marine 
algae have been outlined, and it has been sug- 
gested (2) that such classifications could be 
useful in determining whether certain ecological 
situations are characterized by the same types of 
algae. As such an approach has been used with 
partial success, it might well be expanded to include 
a wider range of conditions and geographical 
areas in order to evaluate its dependability. 

Unfortunately the information relative to the 
chemical and physical factors in an atoll environ- 
ment is far from complete. The information we 
do have indicates that the quantitative differences 
in these factors are probably too small to ac- 
count for distributional patterns among the 
algae and other organisms present. Radiological 
studies to aid in determining the relative amounts 
of minerals about the atoll should be made, and 
these, in conjunction with mineral nutrition 
studies using radioactive tracers, might help to 
clarify the role of minerals in the atoll environ- 
ment. It has been proposed that the life form of 
an alga may be the prime factor in determining 
its distribution, and some studies indicate this 
to be the case. However, sufficient discrepancies 
exist to warrant further study of the problem. 

In summary it can be stated that the occur- 
rence of an alga in a particular ecological niche 
is dependent upon many different factors, among 
which the following must be included: the life 
form of the alga, the chemical and physical factors 
of the environment, the dynamic factors of wave 
action and immersion, and the biological factors 
imposed by other organisms in the vicinity. 


(1) Cloud, P.E., Jr., 1952, Preliminary report on 
geology and marine environment of 

Onotoa Atoll, Gilbert Islands, Nat. Res. 
Counc., Pacific Science Board, Atoll 
Res. Bull., 12:6-73. 

(2) Feldmann, J., 1951, Ecology of Marine 

Algae. In G.M. Smith, Manual of Phy- 
cology, Chronica Botanica Company, 
Waltham, Mass. pp. 312-334. 

(3) Ford, W.L., 1949, Radiological and Salinity 

Relationships in the Water at Bikini 
Atoll, Trans. Amer. Geophvs. Union, 
30(1): 46-54. 

(4) Gilmartin, M., 1956, Personal communica- 


(5) Johnson, J.H., 1954, Fossil Calcareous Algae 

from Bikini Atoll. Bikini and Nearby 
Atolls, Part 4, Paleontology, Geol. 
Survey Prof. Paper 260-M, pp. 537-543. 

(6) Lowman, F.G., Palumbo, R.F., and South, 

D.J., 1957, The Occurrence and Dis- 
tribution of Radioactive Non-fission 
Products in Plants and Animals of the 
Pacific Proving Grounds. Applied 
Fisheries Laboratory, Univ. of Washing- 
ton., U.S. Atomic Energy Commission 
Report UWFL-5L 

(7) Munk, W.H., Ewing, G.C., and Revelle, 

R.R., 1949, Diffusion in Bikini Lagoon. 
Trans. Amer. Geophys. Union, 30 (1): 

(8) Odum, H.T., and Odum, E.P., 1 955, Trophic 

Structure and Productivity of a Wind- 
ward Coral Reef on Eniwetok Atoll, 
Ecol Monogr., 25(3): 291-320. 

(9) Sargent, M.C., and Austin, T.S., 1954, Biolo- 

gic Economy of Coral Reefs. Bikini and 
Nearby Atolls, Part 2, Oceanography 
(Biologic), Geol. Survey Prof. Paper 
260-E, pp. 293-300. 

(10) Taylor, W.R., 1950, Plants of Bikini. Univ. 

of Michigan Press, Ann Arbor, Michi- 
gan, 218 pp. 

(11) Wells, J.W., 1954, Recent Corals of the 

Marshall Islands. Bikini and Nearby 
Atolls, Part 2, Oceanography (Biolo- 
gic), Geol. Survey Prof. Paper 260-1, 
pp. 385-486. 


F.R. FOSBLRO: One must keep in mind that in the Pleis- 
tocene there have been some violent fluctuations with great 
changes in sea level and that the changes are of such an 


extent that one would not think in the same terms. The 
interpretation of a steady state existing since the Miocene 
cannot be rigidly applied. 





Division of Fisheries and Oceanography, Cronulla, Australia. 

The microbiology of coral atolls is a sadly 
neglected field of marine research. We have little 
information, apart from the studies on the 
bacterial precipitation of calcium carbonate made 
by Drew (6), Lipman (7), and Bavendamm (5) ; 
some notes on blue-green algae associated with 
coral reefs by Baas Becking (1 ) ; and some con- 
clusions that may be drawn by inference from the 
work of Baas Becking and Wood (3) and Baas 
Becking, Wood, and Kaplan (4). 

Drew came to the conclusion that large-scale 
precipitation of calcium carbonate was caused in 
tropical waters by a marine bacterium which he 
called Bacterium calcis. He showed that this 
bacterium could reduce nitrates to ammonia, 
which reacted with the bicarbonate in sea water 
to cuase the precipitation of calcium carbonate, 
the calcium being derived from calcium sulphate. 
This is, of course, theoretically possible in a 
region where the calcium and carbonate equilibria 
are in delicate balance near the saturation point 
for calcium carbonate. Lipman questioned 
Drew's hypothesis and considered the production 
of ammonia from organic matter was the impor- 
tant factor in calcium precipitation by marine 
bacteria. He also pointed out that there were 
so few bacteria in the open sea that extensive 
precipitation was unlikely. Bavendamm showed 
that there were enough bacteria in bottom depo- 
sits to cause extensive calcium carbonate preci- 
pitation, but concluded that there were no spec- 
cific bacteria concerned. 

Zobell (9), reviewing the situation, states that 
fewer than 5 per cent of bacterial species in the 
sea are endowed with the ability to liberate free 
nitrogen from nitrate or nitrite in the presence 
of abundant organic matter. My own observa- 
tions show that few marine bacteria can liberate 
either nitrogen or ammonia from nitrates, and 
1 doubt whether this process is important in 
calcium carbonate precipitation. 

Possibly of greater significance in microbial 
calcium carbonate precipitation is the utilization 
of carbon dioxide by autotrophic bacteria and 
photosynthetic microorganisms. There is no 
information on the occurrence or abundance of 
such organisms in the coral reef biocoenosis. 

t Presented in abstract by R.F. Scagel. 

The environmental factors which may affect 
lime deposition include temperature, pressure, 
hydrogen ion concentration, and rcdox potential. 
In fresh water, salt concentrations also have an 
effect, but in the sea, these do not vary sufficiently 
to be important. 

The effect of temperature on calcium carbonate 
precipitation is stressed by Sverdrup ct al. (8), 
who show that, because of the rapid decrease of 
solubility of carbon dioxide with rise of tempera- 
ture, extensive deposition is possible only in 
warmer waters. These authors suggest, too, 
that, as pressure increases the solubility of carbon 
dioxide, lime deposition will be confined to rela- 
tively shallow waters. 

It should be remembered that the carbonate 
equilibrium is very easily reversible, and that 
precipitation presupposes re-solution with a 
slight change of conditions in the right direction. 
Thus, organisms removing carbonate by photo- 
synthesis or photoreduction may, and do, con- 
tinue to produce it in the dark by respiration. 

Baas Becking and Wood, and Baas Becking, 
Wood and Kaplan have considered the relation 
between pH/Eh, along with the microorganisms 
and the environment for a number of biocoen- 
oses. In the first paper, the limitations of a 
number of autotrophs by the environment was 
studied; in the second, the chemical equilibria 
imposing possible limitations were delineated, 
and this has been carried still further by Baas 
Becking (in press). It is unfortunate, but unavoid- 
able, that no observations were made in coral 
reef areas, so we can get only an approximate and 
theoretical picture, of these regions. In the 
estuaries of temperate regions, the pH of the 
water where measured ranged from 7.0 to 9.4, 
with a mean of 8.25, while the muds ranged from 
near 5.0 to near 9.5, with a mean of 7.4. 

The estuarine waters had a mean Eh of +362 
mV with a range from + 150 to + 500 mV, while 
the muds ranged from + 600 to - 350 mV with 
two means, one at about + 325, and the other at 
about - 75 mV. 

Biologically speaking, all autotrophic processes 
except sulphate reduction and photoreduction 



could occur in cstuarine waters, while sulphate 
reduction and photorcduction of carbon dioxide, 
as well as the other autotrophic reactions are 
possible, and can be shown to occur in the estua- 
rine muds. Baas Becking and Wood consider 
that the sulphur cycle dominates the muds, and 
that these in turn dominate the water in shallow 
estuaries. The lower mean Eh of the muds 
reflects this dominance. 

In fresh- water limestone regions studied by 
these authors, the pH ranged from 5.5 to 9.0, with 
a mean of 7. 19, while the Eh ranged from -+ 100 
to +500 mV with a mean at 333 mV. The 
pH shows that bicarbonate ion dominated the 
waters, even those with high concentrations of 
calcium (at 7.2, there will be little COj ion). In 
sea water, at a pH over 8, the equilibrium of the 
CO 2 , HCO 3 , CO 3 would move towards the 
carbonate. At that pH there will be no free 
CO 2 , and 4 parts of bicarbonate to one of 
carbonate ion. A change in the pH to 9.4 will 
reverse the bicarbonate-carbonate ratios to 1 part 
of bicarbonate to about 3 parts of carbonate ion, 
i.e., carbonate will become the limiting factor in 

In temperate estuaries, a pH of 9.3 - 9.4 is 
reached on shallow flats with a large plant popula- 
tion, e.g., of Zostent, in the middle of the day. 
Although, theoretically, calcium carbonate 
should actively precipitate under these conditions, 
the greater acidity of the muds will limit the 
amount actually deposited. Another limit is 
imposed by the short duration of these high pH 
values, and the counter-effect of respiration at 

Coral reefs differ from the estuaries studied by 
Baas Becking and his colleagues in having, on the 
whole, less photosynthctic plants, and conse- 
quently less organic detrital matter in the muds. 
This should lead to a lower activity of the organ- 
isms of the sulphur cycle, which depend on 
heterotrophic bacteria to produce the low Eh 
required for starting sulphate reduction. The 
green and purple sulphur bacteria also require 
low Eh values to initiate the oxidation of sulphy- 
dryl. These regions will also differ from the 
fresh-water limestone regions in having a con- 
siderable amount of calcium sulphate which was 
absent in the areas studied by Baas Becking, 
Wood and Kaplan. 

It can be safely predicted that the alkaline limit 
of the coral reef environment will be controlled 
by carbonate, and the lower Eh limit by the 
amount of organic matter, i.e., reducing material. 


The other limits cannot be estimated except by 
actual observation. 

In temperate estuaries, and in mangrove 
swamps, phosphate is largely controlled by the 
sulphur cycle in the mud (2). The insolubility of 
the sulphides of iron cause the release of phos- 
phoric acid from ferric phosphate in the presence 
of hydrogen sulphide. An alternative method for 
the collection and storage of phosphate by 
microorganisms is the chelation of phosphate in 
the pcctic sheaths, also described by Baas Beck- 
ing and Mackay. I have noticed that encrusting 
growths of blue-green algae usually have a zone 
of sulphate reduction immediately below them, 
and could thus use the sulphur cycle to obtain 
their phosphate. However, one would expect 
the pectic chelation mechanism to be of greater 
importance in the phosphorus cycle on coral 
reefs. Nitrogen fixation may also be an important 
function of the Nostocaceae on coral reefs. 

It will be realized from the account that 1 have 
given that we are largely in a field of pure specula- 
tion when we consider the microbiology of coral 
reefs, but that we have a priori some means of 
approach to the problem. This approach should 
include (1) the study of the microbial compo- 
nents of the coral ecosystem, (2) their interaction 
and cumulative effect on the environment, and 
(3) the actual biological mechanism of lime 
deposition especially as skeletal material. 


(1) Baas Becking, L.G.M., 1951, Notes on some 

Cyanophyceae of the Pacific region, 
Proc. Kon. Ned. Akad. Wet., 54: 2. 

(2) Baas Becking, L.G.M., and Mackey, Mar- 

garet, 1956, Biological processes in the 
estuarine environment, V: The in- 
fluence of Enteromorpha on the environ- 
ment, Proc. Kon. Ned. Akad. Wet., 
B59: 109-123. 

(3) Baas Becking, L.G.M., and Wood, E.J.F., 

1955, Biological processes in the estua- 
rine environment, I &1I: Ecology of 
the sulphur cycle, Proc. Kon. Ned. 
Akad. Wet., B58: 160-181. 

(4) Baas Becking, L.G.M., Wood, E.J.F., and 

Kaplan, I.R., 1957, Biological processes 
in the estuarine environment, X: The 
place of the estuarine environment 
within the aqueous milieu, Proc. Kon. 
Ned. Akad. Wet., B60: 88-102. 


(5) Bavendamm, W., 1932, Die mikrobiolo- (7) Lipman, C.B., 1924, A critical and experi- 

gische Kalkfallung in der tropischen mental study of Drew's bacterial 

See, Arch. Mikrobiol., 3: 205-276. hypothesis in CaCO 3 precipitation in 

the sea, Carnegie lust. Wash. Dept. 

(6) Drew, C.H., 1914, On the precipitation of Mar. #/W., 19: 179-191. 

calcium carbonate in the sea by marine (8) Sverdrup, H.U., Johnson, M.W., and Flem- 

bacteria, and on the action of denitrify- ing, R.H., 1942, The Oceans. Prentice 

ing bacteria in tropical and temperate Hall, N.Y. 

seas, Pap. Tortugas Lab. Wash., 5: (V) Zobell, C.E., 1946, Marine Microbiology. 

7-45. Chronica Botanica Co. Waltham, Mass. 





Auckland University College, Auckland, New Zealand. 

Our present knowledge of the biogeography of 
the South Pacific stems from contributions spread 
over a number of years dealing with algal zona- 
tion and ecology in South America (23,24,25,26), 
Australia (9,1,12,20,31,32), and New Zealand 
(21,8,4,5,2,3,10,11,15). The subantarctic islands 
have also been examined, the first records being 
those of Hooker (14) and Harvey (13), but there 
are recent ones for the New Zealand subantarctic 
islands (6), for the Cro/et Is. (17) and for the 
subantarctic islands generally (18). Another 
island flora that is relevant to any discussion 
on biogeographical provinces is that of Juan 
Fernandez (16). 

The present author believes that proper appre- 
ciation and understanding of the provinces is 
most readily secured by using as a basis the major 
categories recognized by Stephenson (27,28.29) 
in his work on the marine fauna and flora of 
South Africa. The three major categories were: 

(a) A warm water component, predominant 
at Durban and gradually decreasing to Port 
Nolloth. This can be regarded as essentially a 
subtropical province. 

(b) A cold water component associated with 
the cold upwclling on the S.W. African coast, 
and gradually disappearing towards Cape Town. 
This can be regarded as forming essentially a 
cool temperate province. 

(c) A southern intermediate flora of the Cape 
district which can be regarded as a warm tem- 
perate province. 

These major groups were correlated with 
changes in mean sea temperature, and Stephenson 
(29) related them to the main southern littoral 
faunas as described by Eckman based upon 
intervals of 10C. These are as follows: 

( 1 ) Tropical and sub-tropical faunas : minimum 
temperature not below 20C. 

(2) Warm temperate faunas: minimum tem- 
perature 10-20C. 

(3) Cool temperate faunas: minimum tem- 
perature 0- 10C. 

(4) Antarctic faunas: minimum temperature 
0C or lower. 

t Presented in abstract by M.S. Doty. 

A similar ten degree temperature barrier has 
been proposed by Setchell (22) as operative for 
marine algal zones. Recent work by Bennett 
and Pope (1) on Australian faunas suggests that 
the minimum sea temperature boundary of 10C 
between warm and cold temperature faunas may 
need to be raised to 11.5C. In Australia, the 
principal marine faunal and floral provinces 
were first proposed by Hedley but they have been 
amended since then. At the present time the 
following can be recognized: 

(a) The Banksian from Cape York southwards. 

(b) The Solanderian in the Coral Sea area. 

(c) The Peronian in Southern Queensland and 
Northern New South Wales. 

(d) The Maugean in Bass St. and around 

(e) The Flindersian in Victoria and South 

(f) The Dampenan in Western Australia and 
up to the Northern Territories and the 
Gulf of Carpentaria. 

Recent work on the littoral faunas and floras 
has served to substantiate these provinces (1), 
and they have also been shown to be valid for 
the pelagic Dinoflagellates (33). Of these Austra- 
lian provinces, the Maugean must be regarded 
as cool temperate, being associated with cold 
antarctic waters. The Peronian and Flindersian 
provinces are both warm temperate, whilst the 
Solanderian and Damperian are tropical. Fur- 
ther work on those parts of Australia just north 
of the Peronian and Flindersian provinces will 
probably show (hat there are two small subtrop- 
ical provinces characterized by a mixture of 
tropical and warm temperate species. 

In New Zealand, Moore (19) has summarized 
the distribution of some 200 marine algae, and 
as a result she recognized seven distinct provinces 
which can be compared with those of Powell 
for the marine fauna and of Cockayne for the 
terrestrial plants. The present writer, whilst 
agreeing generally with the provinces of Moore 
and Powell is of the opinion that some modi- 
fication is desirable on the basis that has been 
outlined above. The Kermadec algal flora are 



based on old lists (1 ) 9 and a recent collection 
clearly contains tropical, e.g., Caulerpa racemosa, 
C. wekbiana, Pocockiella nigrescens, and warm 
temperate species, and can therefore be regarded 
as a sub-tropical province. Such information 
as is at present available indicates that Norfolk 
Is. falls into the same province. Off the shores 
of South America, the flora of Juan Fernandez 
suggests affinities with Australia and New 
Zealand, but there is a distinct warm water 
element (e.g., Hydroclathrus, Padina, Micro- 
dictyon japonicum, Chaetomorpha antennind) 
which justifies treating it as subtropical. The 
Aupourian or Auckland province is definitely 
warm temperate (11), the proportion of warm 
temperate species gradually decreasing towards 
Cook Strait. The subtropical convergence crosses 
New Zealand in Cook Strait and, as the mini- 
mum sea temperature of the convergence is 
around 10 C C, it forms the logical boundary 
between the warm temperate province of the 
northern island arid the cool temperate province 
of the South Island. The Chatham Islands lie 
very close to the sub-tropical convergence, and 
they possess a flora with warm and cool temperate 
elements. There appears, however, to be a 
slightly higher proportion of warm temperate 
elements, and it may therefore be placed in the 
warm temperate Auckland province, or as a 
sub-province of the Auckland province. 

The marine algal flora of most of the South 
Island of New Zealand must be regarded as cool 
temperate, and it would seem to have a relation- 
ship with the Maugean province of Bass Strait 
and Tasmania. It is to be noted that although 
Tasmania lies just north of the sub-tropical 
convergence nevertheless the waters of the cold 
antarctic current maintain a cool temperate 
flora and fauna. The present writer considers 
that the central and intermediate provinces of 
New Zealand should be merged into a single 
province, the Zealandian. 

The Forsterian (Stewart Is.) and Antipodean 
(= Rossian of Moore) provinces of New Zealand 
comprise the subantarctic islands. Both provinces 
contain a large proportion of southern species, 
so much so that it seems desirable to recognize 
a sub-antarctic province which would also include 
Kerguelen, Crozet, and Heard Islands. In this 
case, therefore, the minimum sea temperature 
boundary would be less than 10C. It may be 
suggested that further work in and around Mag- 
ellan Straits, Tierra del Fuego, and the Faukland 
Islands may show that they also represent a sub- 
antarctic province with a flora differing from that 
of the cool temperate province to the north and 

the extreme antarctic province of the Antarctic 
continent and adjacent Islands. 

There remain the coral reefs and atolls of the 
South Pacific. Our knowledge of the marine 
algal floras of these islands is at present very 
meagre, but it can be suggested that they repre- 
sent essentially a tropical province which can con- 
veniently be called the Central Pacific province. 
It remains to be seen whether New Caledonia, 
the New Hebrides, and the Solomon Islands 
group, now placed in the Solandcrian province 
have a flora sufficiently similar to that of the 
Central Pacific to justify the extension eastwards 
of the Solanderian province. 

Some further confirmation of the conclusions 
that have been reached above can be derived from 
a comparative study of the basic zonations in 
the South Pacific, especially those of Australia 
and New Zealand. Guiler (12) has compared 
(Table 1) the basic (i.e., not greatly exposed nor 
greatly protected) zonation of Tasmania with 
that of New Zealand, Chile, Victoria, and S.W. 
Africa; and he considers that ecologically the 
zonation in Tasmania indicates a greater affinity 
with that found in New Zealand than with that 
of southern Australia. In making the compara- 
tive study, the basis used is that proposed by 
the Stephensons (30). 

In Table 2 a comparison is given of the basic 
zonation of exposed and protected shores in the 
Auckland province of New Zealand with the 
zonation reported from the Zealandian province 
of the S. Island (Banks Peninsula). 

Finally a comparison may be made of the basic 
zonation from three of the Australian provinces 
(Table 3). 

The distinction between the tropical flora and 
the other two is very evident. The similarity 
between the Peronian and the Aucklandian 
provinces is also brought out. In New Zealand, 
Pyura is more frequent in the South Island, 
and it evidently is a representative of the cool 
temperate zone. A fuller comparison of the 
varied zonations that have been described from 
the different provinces is really essential to 
bring out the finer points that demonstrate the 

Whilst our knowledge of the algae and the 
algal zonations in many parts of the South 
Pacific is as yet lacking or very inadequate, 
nevertheless from the data available it would 
appear that the picture that has been drawn 
represents the situation as it is at present. It 
may also serve to highlight those areas from which 
further information would be very valuable. 



Table 1. 


New Zealand 


S.W. Africa 















Elminius or 

Algae or 




Hormosira or 






Ecklonia Cystophora 
Carpophyllum Durvillea 
Lessonia or 

Ecklonia i 


Table 2. 

Hauraki Gulf : Piha 1 Poor Knights Is. Banks Peninsula 
(Protected) (Exposed) (Exposed) (Exposed) 

Littorinids Melaraphe Melaraphe Mclaraphe 
Supra-littoral fringe (Mclaraphe) | 
Myxophyceae Bostrychia Myxophyceae Bostrychia 

i Chamaesipho spp. Chamacsipho Chthalamus Chamaesipho 
Elminius Volsella i Modiolus Elminius Serpulid worm 
Saxostrea Gigartina alveata Apophloea Elminius 
Mid-littoral Corallina- Pachymenia Corallina- Mediolus 
Hormosira Mytilus i Novastoa Mytilus 

Sublittoral fringe Carpophyllum Durvillea (1 sp.) Xiphophora Durvillea (2 sp.) 
Ecklonia Carpophyllum Carpophyllum 

Supra-littoral fringe 

Sub-littoral fringe 

S. Australia 







Table 3. 
I New South Wales 

Tetraclita (exposed) 















(Chthalamus -Tetraclita) 






(1) Bennett, I., and Pope, E.G., 1953, Inter- 

tidal Zonation of the Exposed Rocky 
Shores of Victoria, together with a 
re-arrangement of the biogeographical 
provinces of temperate Australian 
shores, Aust. Journ. Mar. and F. W. Res., 
4(1): 105. 

(2) Beveridge, W.A., and Chapman, V.J., 1950, 

The Zonation of Marine Algae at Piha, 
New Zealand, in relation to the tidal 
factor (Studies in Inter-tidal Zonation 
2), Pac. Sc/., 4(3): 188. 

(3) Carnahan, J.A., 1952, Inter-tidal Zonation 

at Rangitoto Island, New Zealand 
(Studies in Inter-tidal Zonation 4), Pac. 
Sci., 7(1): 35. 

(4) Chapman, V.J., 1950, The Marine Algal 

Communities of Stanmore Bay, New 
Zealand (Studies in Inter-tidal Zonation 
1), Pac. SW., 4(1): 63. 

(5) 1956a, A Revision of the 

Marine Algae of New Zealand, Pt, 
I: Myxophyceae and Chlorophyceae, 
Journ. Linn. Soc. Bot., 55: 333. 

(6) , 1956b, Marine Algal 

Provinces and Zones in New Zealand: 
their place in a World System, Proc. 
1st Geog. Conf., N.Z. Geog. Soc., p. 27. 

(7) Cotton, A.D., 1911, Marine Algae from 

the North of New Zealand and the 
Kermadecs, Kew Bull., 61: 253. 

(8) Cranwell, L.M., and Moore, L.B., 1938, 

Inter-tidal communities of the Poor 
Knights Islands, N.Z., Trans. Rov. 
Soc. N.Z., 67: 375. 

(9) Dahin, W.J., Bennett, I., and Pope, E.C., 

1948, A Study of Certain Aspects of the 
Ecology of the Inter-tidal Zone of the 
New South Wales Coast, Aust. Journ. 
Sci. Res. Ser. B., 1 (2): 176. 

(W) Dellow, U.V., 1950, Inter-tidal Ecology at 
Narrow Neck Reef, New Zealand (Stu- 
dies in Inter-tidal Zonation 3), Pac. 
Sci., 4(4): 355. 

(11) , 1955, Marine Algal Ecol- 
ogy of the Hauraki Gulf, New Zealand, 
Trans. Roy. Soc. N.Z., 83 (1): 1. 

(12) Guiler, E.R., 1952, The Nature of Inter- 
tidal Zonation in Tasmania, Pap. Proc. 
Roy. Soc. Tasm., 86: 31. Handbook of 
the New Zealand Flora, London. 

(13) Harvey, W.H., 1855, Algae in J.D. Hooker, 

Flora Novae Zelandiae, 2:211, London. 

(14) Hooker, J.D., 1847, Flora Antarctica, 2, 


(15) Knox, G.A., 1953, The Inter-tidal Ecology 

of Taylor's Mistake, Banks Peninsula, 
Trans. Roy. Soc. N.Z., 81 (2): 189. 

(16) Levring, T., 1941, Die Meeresalgen der 

Juan Fernandez-Inseln, Nat. Hist. 
Juan Fern\ and Easter Is., Ed. C. Skott- 
sberg,2: 601. 

(17) ___ , 1944, Meeresalgen von 

den Crozet Inseln und Kerguelen, Ark. 
Dot., 31 (8): 1. 

(18) _______ - , 1945, Marine Algae 

from some Antarctic and Subantarctic 
Islands, Lunds. Univ. Arss. N.F. Avd. 2, 

(19) Moore, L.B., 1949, The Marine Algal Pro- 

vinces of New Zealand, Rov. Soc. N.Z. 
Proc. 6th Sci. Congr., p. 187. 

(20) Newton, L., and Cribb, A.B., 1951, Some 

Aspects of Algal Ecology in Britain and 
Australia, Research, 4: 449. 

(21) Oliver, W.R.B., 1923, Marine Littoral Plant 

and Animal Communities in New Zea- 
land, Trans, and Proc, N.Z. Inst., 54: 496. 

(22) Setchell, W.A., 1920, Stenothermy and 

Zone Invasion, Amer. Nat., 54: 385. 

(23) Skottsberg, C., 1907, Zur Kenntnis der Sub- 

artanktischen und Antarktischen Meere- 
salgen I. Phaeophyceen, Wiss. Ergebn. 
d. Schwed. Sildpolar Expd. 1901-03, 
4, Stockholm. 

(24) . ________ ..,1921, Marine Algae I: 

Phaeophyceae (Botanische Ergebnisse 
der Schwediochen Expedition nuch 
Patagonien unde der Feuerland, 1907- 
09), K. Svenska Vet-Akad. Hand, 61, 

(25) __ ._. _, 1923, Marine Algae II: 

Rhodophyceae, Ibid., 63, Stockholm. 

(26) , 1950, Sodra Ishavets 

Algflora, K. Svenska Vetenskap-Akad- 
Arsbok, p. 367. 

(27) Stephenson, T.A., 1939, The Constitution 

of the Inter-tidal Fauna and Flora of 
South Africa, Part I, Journ. Linn. Soc. 
Zoo I., 40: 487. 

(28) , 1944, Ibid., Part II, Ann. 

Natal Mus., 10: 261. 

(29) , 1948, Ibid., Part III, Ann. 

Natal Mm., 11 (2): 207. 



(30) Stephenson, T.A. and Stephenson, A., 1949, of the algal ecology, Trans. Roy. Soc. 

The Universal Features of Zonation be- S. Aust., 71 (2): 228-229. 

tween Tide Marks on Rocky Coasts, (32) , 1948, Ibid., II: ThePen- 

Journ. EcoL, 37 (2): 289. nington Bay Region, Ibid., 72 (1): 144. 

(33) Wood, E.J.F., 1954, Dinoflagellates in the 

(31) WomersIey,H.B.S., 1947, The Marine Algae Australian Region, Aust. Journ. Mar. 

of Kangaroo Is. I: A general account and F. W. Res., 5 (2): 171. 







Deputy Director, VInstitut Oceanographique, Nhatrang, Vietnam. 

The Nhatrang bay, and the sea coast of Central 
Vietnam in general have a physiognomy which 
distinguishes them from the rest of the whole 
coast from the ecological point of view. 

The "Annamitic Cordillera" ends there with 
rocky outgrowths delimiting sandy bays while 
alluvial deposits brought in by large rivers con- 
tinue to build elsewhere muddy and sandy coasts. 
Climatologically, the coast of Central Vietnam 
owes the Cordillera a climate diametrically op- 
posed to that of the rest of the country. When 
in the North and the South the summer monsoon 
brings in the rain, the dry season (from February 
to August) prevails in the Centre. The yearly 
average of rains amounts to 1,441 mm against 
1,979 in Saigon. 

At Nhatrang, the thermic regime is also the 
same as in Saigon, with a small temperature 
difference (4) between the warmest and the 
coolest month. 

Finally, oceanographically the central coast is 
distinct. There, in fact, diurnal and semi-diurnal 
tides manifest themselves on an equal basis. The 
tide regime is a mixed one as compared with 
a semi-diurnal regime in the South and a diurnal 
regime in the North. (We are awaiting details 
about the harbour establishment along the Viet- 
nam coast). 

Thus the physical, climatological, and oceanog- 
raphical data put emphasis on the interest of 
a comparative study of the ecology of the central 
sea-coast and that of the rest of Vietnam. 

We began with the study of the submarine 
vegetation at Nhatrang, mainly on rocky spots. 
We took, for example, the rhyolitic point of 
Qiuda. We could distinguish the three general 
bionomical zones of Stephenson (1949), viz. 
a Littorine zone, a Cirripede zone, and the sub- 
littoral fringe. But the zones which can be easily 
evidenced are from top to bottom: 

(a) the Calothrix pilosa Born, and Flash girdle, 
30 cm wide, watered by high sea spray (embruns) 
and submerged under the biggest spring tides 
(lunar tropic). This is a discontinued girdle, 
confined to rough substrata. 

Upward accessories: Littorina scabra Lin., 

t Presented by M.S. Doty. 

Tectarus gramdaris Gray, Lygia sp., ubiquists: 
Grapsus grapsus. 

(b) the Brachytrichia maculans Com. girdle, 
covering 80% to 90% of the area. Another typical 
growth, Littorina scraba Lin., has a maximum 
thickness which changes according to the tide. 

In the anfractuosities are the Isognomon sp. 
and Pollicipes mittellae ("mode battu"). And at 
the lower limit are the patella: Cellana amussilata 
Reev., Acmaea saccharina Linn. A. granostriata 
Sch Patella aster Reev., etc. 

(c) the Ostrea cucullata girdle, situated at the 
mid-tide level with Ectocarpus braeviarticulatus 
J.Ag., Chnoospora pacifica J.Ag., Ge Helium pusil- 
lum Le Jolis ("mode battif), Acrocystis nana 
Zan. On the walls, eurytopics colonies of Paly- 
thao sp. 

(d) the Gelicliella acerosa girdle at the low water 
levels is the large Phaophyccae zone : Sargasswn 
ssp. Turbinaria ornata and Turbinaria sp. ("mode 
battu") Padina commersoni Bory., Chnoospora 
implexa J. Ag. ("mode peu battu a calme"). The 
great phaeophyceae shelter abundant epiphytic 
flora. On padina commersoni Bory, for example, 
we found Griffithsia metcalfii Tseng, G. tennis 
C. Ag.,Acrochaetumgmci/e, Centrocerasclavatum 
Mont., Ceramium mazatlanense Daws., Poly- 
siphonia sp. Great acorn-shells (Balanus tintin- 
nabulum) were covered with Melobeseae or Ralfsia 
sp. at the beaten spots; on the less beaten rocks 
were Gastrochaena cymbium Sprengl. On the 
scattered rocks you find Enteromorpha intestinalis 
Link. On the small grounds where the waves 
are not sharp are the tufts of Ectocarpus mitchellae 
mixed with Enteromorpha clatharata J. Ag. and 
E. Kyllini Blid. 

Inside the beach rock blocks we found 
Lithodomus lima Jouss., L. Malaccarnus Reev., 
Pholadidea far rot i Jouss., Area turella, several 
crustaceous: Gonodactylus, Alpheideae, Isopoda; 
Spongiaires and Polychaeta. 

(e) the infra-littoral level begins with Thalassia 
Hemprichii Asch., Halophila ovalis Hook, Diplau- 
thera Uninervis Asch. that is found only at the low 
spring waters. You find there Colpomenia sinuosa 
Derb & Sol., Padina commersoni, Gracilaria 



crassa Harv., Bornetclla oligospora, Neomeris dcma setosa and other sea-urchins, Mulleria sp., 

annulata Dick, Halimeda gracilis Harv., Struvea the Crinoids. It is also the coral zone whose 

anastomasans Pice., Galaxaura fastigiata Dec., study is being conducted by M.M.G. Ranson 

etc At the lower part, you see between rocks: and Nguyen-Thanh-Tri. 

Ceratodictyon spongiosum Zan., Halymenia dila- We do not yet have data about the deep lower 

tata Zan., Liagora ceranoides Lam., Liagora infra-littoral zone. Dawson (1954) found Gala- 

farinosa Lam., Q. Jilamentosa Chou, etc..., Dia- xaura Vietnamiensis Daws, at 30 m deep. 


M.S. DOT Y: Is the 4 C. temperature difference referred P.H. HO: The temperatures given are for air. 
to for air or water? 





University of British Columbia , Vancouver, Canada, 

In comparison with the progress in our knowl- 
edge of most groups of plants especially con- 
cerning their life-histories and distributions 
the advances made in marine phycology have 
been relatively slow. The limited access to living 
material or to the facilities to maintain the larger 
marine algae in the living condition for a pro- 
longed period of time, the difliculties of col- 
lection particularly in the subtidal zone and 
the lack of any extensive direct economic value 
until recent years have all contributed to this 
slow progress. However, in spite of these dif- 
ficulties, there has been a considerable amount 
of interest in the marine algae, including many 
studies of their ecology. Although this interest 
has been fairly widespread in a number of 
countries of the world, until recently there has 
been little activity in the field of marine benthic 
algal ecology on the Pacific Coast of North 
America and nothing of a comprehensive nature 
has been published for this area. It is an ana- 
chronism that this should be so in a region which 
received such prominence some fifty years ago 
through the efforts of a pioneer in the field, the 
late William Albert Setchell. 

Although the knowledge of the effect of tem- 
perature on the world-wide distribution of plants 
both horizontally and vertically had developed 
gradually over a period of many years, it was 
only during the last hundred years that the atten- 
tion of phycologists was brought to a consider- 
ation of the reasons for the observed distributions 
of the marine algae. The historical development 
of this trend of thought and investigation during 
this early period has been reviewed by Setchell 
(30). Starting over fifty years ago and extending 
through a series of papers from 1893 to 1935, 
Setchell made a noteworthy attempt to explain 
the world-wide distribution of marine algae, and 
especially members of the Laminariales on the 
Pacific Coast of North America, on the basis of 
latitudinal and seasonal temperature distributions. 
The physical data available during this early 
period were limited, but many of the principles 
set forth by Setchell concerning the distributions 
of marine algae are as sound now as when they 
were first proposed. Except for more precise 
knowledge of the physical and chemical factors 

of the environment and the distributions of the 
algae concerned, much of SetchelFs ecological 
work can still be used as a good foundation for 
further study. Although it was largely a two- 
dimensional approach to the marine environment, 
Setchell's work made a significant contribution 
to the development of marine algal ecology. 

Lamouroux (23, 24) had suggested the possibi- 
lity that temperature stratification in the sea 
might account for the vertical distribution of the 
marine algae and had considered the effect of 
tides on intertidal zonations, but this trend to 
analyze the vertical distribution of the marine 
algae was not taken up in detail until much later. 
Coleman (12) was one of the first to emphasize 
the use of tide levels to account for the vertical 
distribution of the marine algae in the intertidal 
zone. In a study in Oregon, on the Pacific Coast 
of the United States, Doty (15) has given further 
evidence for the relationship between the verti- 
cal distributions of marine algae and critical tide 

There have been a number of lists of marine 
algae published, and attempts have been made 
not only to relate the floras of one area to another, 
such as that by Okamura (25, 26) in the North 
Pacific, but also to account in a general way for 
distributions on the basis of ocean currents, such 
as that by Isaac (22) in the area around South 
Africa. However, there soon followed a decided 
shift to intertidal studies of regional areas, such 
as that by Feldmann (17) in the Mediterranean 
and Chapman (11) and his students in New Zea- 
land. Some attempts have also been made to 
describe universal features of intertidal zonation 
throughout the world (32). At the same time, 
there has been a tendency to place greater em- 
phasis on the interrelationships between the 
various organisms. 

Many of these intertidal studies have been of 
great value as an initial descriptive stage of in- 
vestigation, and there is a need for further de- 
scriptive studies of this type in new and unde- 
scribed regions. However, the variety of systems 
of nomenclature and terms that have been pro- 
posed by marine ecologists to describe zonation, 
associations, and other ecological concepts have 



frequently only complicated the descriptive study 
rather than succeeded in explaining the observed 
phenomena. This has led to some confusion in 
terminology. It is a debatable point whether there 
can be such a thing as a universal system of classi- 
fication beyond a generalized scheme, such as 
that proposed by Ekman (16), and it is question- 
able whether some of the systems proposed can 
contribute further to progress in marine algal 
ecology even in regional studies without simpli- 
fication or clarification. 

Although the shift in emphasis to the inter- 
relationship of organisms was an important one, 
in some instances this approach has been respon- 
sible for excluding adequate concurrent studies 
of the physical and chemical aspects of the en- 
vironment. It is for this reason that a case may 
be made for reassessing our position in marine 
algal ecology, and a critical evaluation of the next 
steps to be taken to further our progress is timely. 
Perhaps what may be called a three-dimensional 
or an oceanographic approach can be used to 
analyze more precisely various factors in the 
marine environment and the relationship of these 
factors to bcnthic algal productivity. A step in 
this direction has been considered recently with 
some measure of success by Dawson (13, 14) in 
Baja California, Doty (15) in Oregon and Wo- 
mersley (34) in Australia. 

To the oceanographer, the most complicated 
physical or chemical situation to explain may be 
the smallest unit of the environment with which 
he is faced. This is partly a problem of instru- 
mentation. However, it is usually much easier 
not only to recognize significant discontinuity in 
properties, such as temperature, salinity even 
plankton distributions over extensive areas of 
the ocean than in restricted or local regions, but 
also to use such information in describing dyna- 
mic processes. Hence, it is suggested that more 
attention should be given to studies of the general 
distribution of various physical and chemical 
properties in the marine environment in an 
attempt to set up some workable hypotheses to 
account for observed distributions of marine 
algae. In this way we may hope to explain and 
account for biological phenomena rather than be 
satisfied by a description of the phenomena or by 
terms to describe them which do nothing more 
than give names to dynamic aspects of marine 
ecology much in need of logical explanation. 
With increased activity recently in oceanography 
in the Pacific, we may now hope for more abun- 
dant and usable data on some of the more general 
oceanographic properties of the North Pacific. 
In specific cases, and particularly in more 

restricted areas, the ecologist will be forced to 
turn more attention to obtaining in situ physical 
and chemical data before further progress can 
be made. 

One can arbitrarily start by summarizing all 
the factors in the marine environment as geolog- 
ical, physical, chemical, and biological. The 
way in which these are considered may be some- 
what a matter of interpretation. Salinity, for 
example, may be considered directly, as a chemical 
factor, or indirectly as a physical factor respon- 
sible for changes in density and thus contributing 
to the pattern of circulation. Likewise, the nature 
of the substratum may be considered indirectly as 
a geological factor, or directly as a physical or 
mechanical factor restricting or permitting estab- 
lishment of benthonic organisms because of 
particle size. There has been much written on 
many of these aspects of ecological study in 
special cases, but it is suggested that, in a general 
overall reassessment of the environment, an 
attempt be made to proceed from this more 
general position to the particulate. This approach 
may initially lead only to the erection of further 
hypotheses, as the indirect or direct nature of the 
action of any particular factor in the environment 
may ultimately be established only by further 
experimental work either in the field or in the 
laboratory. This is the approach that is being 
promoted in the studies on marine algae being 
carried out on the Coast of British Columbia, 
Canada, some aspects of which will be mentioned 
briefly in this review. 

Although on occasion in the past the position 
of the taxonomist has been questioned in the 
scientific field, it is apparent that the need for 
fundamentally sound taxonomic studies of the 
marine algae is almost as great now as it has ever 
been, particularly since the recent progress in 
knowledge of the fields of biochemistry, cytology, 
and genetics. However, in the sense implied here, 
the work of taxonomist has reached a logical 
conclusion only when it is applied, and it is this 
application that is frequently left for the ecolo- 
gist. As a step in the direction of increasing our 
knowledge of the taxa which comprise the tools of 
the algal ecologist and of completing this de- 
scriptive phase of the study of the marine ben- 
thonic algae, an annotated check list has just been 
completed for the Coast of British Columbia and 
Northern Washington (28). Based on this list, 
further studies are now in progress to augment 
the existing data on the marine flora of British 
Columbia, not only insofar as distributions are 
concerned, but also relating to life-histories, 
growth, reproduction, and seasonal aspects. 

These problems are now being tackled in marine 
laboratories which have running seawater avail- 
able, as well as in the controlled-environment 
tanks and rooms on the campus of the University 
of British Columbia. These fundamental studies 
are basic to all other aspects of ecological 
research, and more especially when an attempt 
is made to use specific organisms as indicators 
of oceanographic conditions. 

Not only does this descriptive phase require 
an adequate consideration of the taxonomic 
aspects, but a more complete description of the 
other factors in the environment is needed. With 
increased activity recently and currently in general 
oceanographic studies of the North Pacific by 
a number of institutions on the Pacific Coast of 
Canada, in Japan, and in the United States, there 
has resulted a considerable body of knowledge, 
but it is still far from adequate for the ecologist, 
particularly for one interested in coastal dyna- 
mics. Circulation problems, temperature, sali- 
nity, oxygen, and in some areas, phosphate and 
nitrate distributions are fairly well known in a 
broad and general sense, but at the present time 
very few restricted areas are known oceanograph- 
ically in sufficient detail. More precise physical 
and chemical data, even in these more restricted 
areas, are also inadequate. The distribution of 
other chemical constituents, and to a large extent 
even a knowledge of the plankton composition, 
distribution, and activity, is almost completely 
lacking. It is obvious that the more data that 
become available the more clearly one can tackle 
problems relating to specific distributions and 
set up field and laboratory studies to test hy- 

It is already apparent that much can be done 
experimentally both in the field and in the labora- 
tory with benthonic marine algae. Many of the 
problems encountered by the ecologist dealing 
with the larger marine algae present unique 
culture problems and require an entirely different 
approach, both in the field as well as in the labo- 
ratory. Some studies on growth and reproduction, 
particularly of some of the larger Laminariales, 
have been done in this region, both in the field 
(27) as well as in the laboratory. Although the 
size of many of the cold-water marine algae add 
new problems, at least some of the stages can be 
carried out to the point where transplants can be 
made from the laboratory into the sea for further 
study. Transplant experiments of natural popu- 
lations of these larger marine algae, even in the 
case of Macrocystis, are quite feasible. 

The successful use of the experimental approach 
in the laboratory is primarily dependent on having 


facilities for maintaining temperature and light 
control, although the size of plants may again 
present certain special problems. Cultures of 
Laminariales have been maintained in con- 
trolled-environment tanks at the University of 
British Columbia for as long as a year, during 
which the complete sexual generations were 
cultured and the young sporophytes reached a 
length of fourteen inches, well past the stage 
where secondary morphological characteristics 
had developed. These studies have permitted 
indisputable identification of the sporophytes to 
genus and in some cases to species. The study of 
cultures in this group suggests that much of the 
early work on gametophytes in the Laminariales, 
and in fact even on the early sporophytes, may be 
in some question. In most of the early studies, 
reported plants were not grown long enough to 
establish beyond doubt the characteristic 
secondary morphological features of the sporo- 
phytes of the genera from which zoospores were 
initially obtained. In the presence of contaminat- 
ing zoospores of other species, there is not other 
way of establishing that the same species or even 
the same genus in the Laminariales was obtained 
in the sporophyte generation succeeding the 
gametophyte generations in culture. 

It would be remiss not to mention much of the 
worthwhile physiological work that has been 
done on marine algae. However, there is a need 
for a great deal more physiological work, parti- 
cularly of the type done by Gail (18, 19,20) in an 
attempt to related physiological processes more 
specifically and directly to the environment and 
ecological problems encountered in the field. 
In physiological studies, there is frequently a 
tendency to proceed more and more deeply into 
special aspects of the physiological behaviour or 
biochemistry of an organism under artificial 
conditions, rather than to project back to the 
field and attempt to explain behaviour under the 
conditions existing in the natural environment. 

Almost all of the quantitative aspects of benthic 
algal productivity in this area have related to 
species of economic interest (27,21). These studies 
have dealt largely with quantities and distribu- 
tion and have contributed little to an evaluation 
or an explanation, in terms of oceanographic 
factors, of the causes for this production. 

Obviously the ideal of a functional interpre- 
tation is dependent on an adequate and balanced 
knowledge of all of the foregoing aspects of 
the qualitative and quantitative features of both 
the organisms and the environment. Many, 
much-needed, data are still lacking. An attempt 
to follow this line of investigation has been started 



on the Coast of British Columbia, and in a some- 
what more restricted area at the north end of 
Vancouver Island in Queen Charlotte Strait. 
Further detailed work is in progress in the Strait 
of Juan de Fuca at the south end of Vancouver 
Island between Vancouver Island and Northern 

The Pacific Coast of Canada is ideally suited to 
a study of benthonic organisms and the effect of 
oceanographic factors on their distribution both 
in the intertidal and the subtidal zones. Although 
the coast of British Columbia is only about 600 
miles long, proceeding directly from the Strait 
of Juan de Fuca to Dixon Entrance, if all its 
various ramifications are included, there is a 
coastline estimated at about 25,000 miles in 
length. The tidal amplitude in this region is 
great, ranging from about 1 1 feet at the southern 
boundary to nearly 26 feet at the northern boun- 
dary. As a result of thorough mixing in the 
coastal region, the upper zone in this area, 
except for a few local anomalies, is characteristi- 
cally rather uniform in temperature at any one 
period and fluctuations occur within narrow 
limits. The annual range in temperature of the 
sea water near the surface is from about 6 degrees 
C to 18 degrees C. On the other hand, because 
of the excessive run-off from large rivers, 
especially through the long mainland inlets, 
there are conditions ranging from practically 
fresh water at one extreme to full ocean salinity 
of about 34% at the other extreme. The oceanog- 
raphic conditions characteristic of the Coast 
thus provide a particularly ideal area in which 
to study the distribution of marine benthonic 
organisms in relation to salinity over a rather 
extensive geographic area. 

Throughout the Coast, the physical nature of 
the substratum ranging from mud and sand 
at one extreme to solid rock at the other deter- 
mines to a large extent the organisms which are 
found in a specific area. However, a comparison 
of the flora and fauna on various types of bottom 
is possible in a number of regions which are 
otherwise oceanographically rather similar. This 
permits a correlation of the distribution of a wide 
variety of plants and animals with other physical 
and chemical factors of the environment. 

Biological observations, extending over the 
whole length of coastline, indicate that there is 
a high degree of uniformity in the populations of 
many benthonic plants and animals extending 
from the Strait of Juan de Fuca to Dixon 
Entrance. This would be expected under the 
relatively uniform conditions of temperature 

indicated. In attempting to correlate the dis- 
tribution of some of these organisms with salinity 
characteristics, as well as other oceanographic 
factors, there are several areas on the Coast 
which could be used for purposes of this study. 
Although some supporting observations have 
been made in the Strait of Juan de Fuca and 
Dixon Entrance, and further work is being done 
in the former area at the present time, the present 
review is restricted largely to a consideration of 
the vicinity of Queen Charlotte Strait near the 
north end of Vancouver Island. 

Although soundings are still somewhat incom- 
plete for the area, a general study of the bottom 
topography in Queen Charlotte Strait indicates 
extensive shallows, particularly along the Van- 
couver Island side of the Strait and around Mal- 
colm Island. In this region an abundant and 
varied intertidal and subtidal flora and fauna 
are supported. In the central part of the Strait, 
and between Nigei and Vancouver Islands, there 
are deeper channels exceeding 100 fathoms in 
depth. These channels are not continuous, 
however, with the deeper waters of the mainland 
inlets and Johnstone Strait, and exhibit physical 
and chemical properties quite distinct from the 

The salinity distribution near the surface in 
Queen Charlotte Strait indicates a general cir- 
culation in a counter-clockwise fashion. The 
run-off from the mainland inlets along the north 
shore and at the east end of the Strait, particu- 
larly from Knight Inlet at the east end of the 
Strait, contributes large volumes of fresh water 
which tends to move seaward at the surface mix- 
ing as it progresses along the north shore into 
Queen Charlotte Sound with the deeper more 
saline water below. The more saline water from 
the open ocean and Queen Charlotte Sound 
moves into the Strait centrally as well as along the 
north side of Malcolm Island. The intrusion of 
high salinity water along the deep channels in 
the central part of the Strait is also apparent. 
This general pattern of salinity distribution, with 
fluctuations to varying degrees near the surface, 
is pronounced in the upper zone to a depth of 
at least 20 meters. 

An analysis of surface salinity data over a 
ten-year period from Pine Island Lighthouse 
(/ through 10), which is near the entrance to 
Queen Charlotte Strait, indicates a salinity 
maximum of about 34% and a minimum of about 
28% with an annual mean of 31.75%. Although 
this station is not fully characteristic of the whole 
of Queen Charlotte Strait, the data available give 



a close approximation of the annual salinity 
fluctuations in the more oceanic part of the 
Strait. Insufficient data are available from 
Pulteney Point on Malcolm Island as yet to 
analyze the seasonal fluctuations in the central 
region of the Strait, but, for the period available, 
a range of about 28% to 32% with an annual 
mean of 28.95% is indicated. 

A comparison of the temperature-salinity 
characteristics of various parts of the Strait and 
the connecting bodies of water by means of T-S 
diagrams indicates the discreteness of the water 
masses typical of Johnstone Strait, Knight Inlet, 
and Queen Charlotte Sound. The T-S diagrams 
for Queen Charlotte Strait indicate a character- 
istically intermediate condition between these 
extremes in properties of temperature and salinity 
for the greater part of the Strait. 

An analysis of surface temperature data over 
a ten-year period from Pine Island Lighthouse 
(7 through JO) indicates a temperature maximum 
of about 12 degrees C and a minimum of about 
5 degrees C, with an annual mean of 8.6 degrees 
C. Although this station, as already indicated, 
is not characteristic of the Strait in all regions, 
it probably gives a reasonable approximation of 
the annual temperature fluctuations for the outer 
part of the Strait. As is true in regard to salinity, 
insufficient data are available from Pulteney 
Point on Malcolm Island as yet to analyze the 
seasonal fluctuations in the central region of the 
Strait, but for the period available a range of 
from about 5 degrees C to 1 1 degrees C with an 
annual mean of 8.25 degrees C is indicated. 

The tidal amplitude in this area is about 17 
feet, so that there is exposed an extensive inter- 
tidal flora and fauna during lowtide periods. 
Although no attempt will be made to analyze the 
vertical distribution of the organisms of the 
intertidal zone here, it may be pointed out that 
many of them can be related to certain tide levels. 

Biological observations have been made 
throughout Queen Charlotte Strait, although a 
more intensive study has been undertaken at 
Hope, Deer, and Malcolm Islands. The observa- 
tions have been restricted so far to the more 
conspicuous algae and invertebrate animals in 
the intertidal and immediately accessible subtidal 
zones. These localities present a transition from 
Hope Island, where the highest salinities are 
encountered, to the north and east sides of the 
Strait where the lowest salinities are found. Deer 
Island and Malcolm Island are intermediate 
between these extremes. 

Some organisms in the area are more cos- 
mopolitan in their distribution, particularly in 
their tolerance to extreme dilution. Extending 
throughout the Strait are forms such as Alaria 
tenuifolia Setchell f. tenuifolia, Cymathere tripli- 
cata (P. and R.) J.Ag., Costaria costata (Turn.) 
Saunders, Costaria mertensii J.Ag., Laminaria 
saccharina (Linnaeus) Lamouroux f. saccharina, 
Nereocystis luetkeana P. and R., Porphyra per- 
forata}. Ag. f. perforata, Rhodomela larix (Turn.) 
C. Ag., Odonthaliafloccosa (Esper) Falk., Mytilus 
edulis Linnaeus, Littorina planaxis Philippi and 
Strongylocentrotus drobachiensis (Miiller). 

Restricted to the region of highest salinity, as 
at Hope Island, are Postclsia palmaeformis Rupr., 
Lessoniopsis littoralis (Farl. and Setch.) Reinke, 
Laminaria setchellii Silva, Pelvetiopsis limitata 
(Setchell) Gardner f. limitata, Dilsea californica 
(J.Ag.) O. Kuntze, Erythrophyllum delesserioides 
J.Ag., Jridaca Uneare (S. and G.) Kylin, Hymenena 
setchellii Gardner, Ptilota asplenioides (Esper) 
C.Ag., Ptilota californica Rupr., Ptilota hypnoides 
Harvey, Mitel/a polymerus (Sowerby) and 
Flustrella corniculata (Smitt). Pleurophycus 
gardneri Setch. and Saund., Pterygophora califor- 
nica Rupr., two forms of Hedophyllum sessile 
(C.Ag.) Setch. and Styela montereyensis Dall are 
also present in the regions of highest salinity but 
are somewhat less restricted in their distribution 
and extend into the Strait almost as far as Deer 

A few organisms extend still further into the 
Strait but only slightly beyond Deer Island. 
Among these are Alaria nana Schrader and Alaria 
marginata P. and R. Still others extend from 
Hope Island past Deer Island down into the 
Strait as far as Malcolm Island, but not as far as 
the east and north sides of the Strait. Among 
these are Macrocystis integrifolia Bory, Egregia 
menziesii (Turner) Aresch. subsp. menziesii, 
Alaria valida (Kjell. and Setch.) f. valida, the 
bullate form of Hedophyllum sessile (C.Ag.) 
Setch., Constantinea simplex Setch., Mytilus 
californianus Conrad, Strongylocentrotus purpu- 
ratus (Stimpson) and Haliotis kamtchatica Jones. 
Isolated populations of the abalone (Haliotis 
kamtchatica), which have been noted further 
eastward in Johnstone Strait, and which may be 
related to local oceanographic features, such as 
upwelling, present somewhat of an anomaly to 
the general distribution. A further exception to 
the distribution described in this group is that of 
Macrocystis integrifolia. Although Macrocystis 
occurs in regions of high or relatively high sali- 
nities and extends as far down the Strait as 



Malcolm Island and over to Numas Island, it 
does not occur in the most exposed areas on the 
open coast where Postelsia and Mitella are en- 

There is a distribution of organisms here which 
follows closely the pattern of salinity distribution 
in the Strait, which in turn reflects the circulation 
within the area. It would be premature to say that 
salinity is directly responsible for the observed 
distributions of all the organisms encountered. 
But one may say that the distribution reflects the 
dependence on waters characteristic of the open 
ocean and exposure. In some instances, it may be 
directly salinity that is a causal factor. The open 
coast, on the other hand, has organisms associated 
with surf conditions. It has been suggested that 
the high oxygen requirement of certain organisms 
is met only in such an exposed environment. 
However, the measurement of oxygen values in 
the sea in this area, does not directly support 
this argument, as the oxygen content of the waters 
within the sheltered Strait is as high or higher 
than in the surf in the exposed regions. This is 
particularly true in the central part of the Strait 
when there is a heavy bloom of phytoplankton 
in the area, at which time the water may be super- 
saturated to as much as 175 per cent. Likewise, 
although it is known that many marine algae 
have a high inorganic phosphate requirement, 
there is no evidence that this nutrient is ever 
limiting in this area within the zone occupied by 
the benthonic marine algae. There is a great 
need for further knowledge of the presence, dis- 
tribution, amounts, and availability of many 
more dissolved inorganic and particulate organic 
substances. The evidence so far points also to 
the need for a study of quantitative removal and 
the rate of removal of such substances and precise 
requirements for growth and reproduction of the 
benthonic marine algae. 

The restriction of certain organisms to surf 
conditions suggests that constant movement' of 
water is required to provide nutrients and gases 
which may be rapidly exhausted from the im- 
mediate micro-environment of the individual 
alga, or in the case of the sessile marine inverte- 
brate, such as Mitella polymerus, to provide 
particulate food. It may be that lowering the 
concentration or removal by dilution or water 
movement of some substances which accumulate 
above a certain level of concentration in the 
micro-environment is just as significant as the 
availability of others. 

In summary, a detailed study of the distribution 
of marine benthonic organisms in Queen 


Charlotte Strait has been limited so far to the 
more conspicuous algae and invertebrates en- 
countered. The relationship of these distributions 
to the salinity distribution indicates that more 
intensive study of the flora and fauna in this area, 
as well as elsewhere on the Coast, will provide 
further supporting evidence of the effect of ocean- 
ographic variables on the distribution of marine 
benthonic organisms and the possibility of using 
such organisms as indicators of oceanographic 
conditions both in time and space. It is hoped 
that this oceanographic approach both quali- 
tatively and quantitatively may then lead not 
only to a clearer understanding and explanation 
of the fundamental relationships between the 
organisms and their environment, but also to an 
understanding of the interrelationships and in- 
teraction among the organisms themselves. 


(1) Anon, 1944, Observations of Seawater Tem- 

perature and Salinity on the Pacific 
Coast of Canada, Fisheries Research 
Board of Canada, Nanaimo; Pacific 
Oceanographic Group, Vol. 5, 1942 
and 1943. 

(2) - - , 1946, Ibid., 6, 1944 and 1945. 

(3) , 1948, Ibid., 7, 1946 and 1947. 

(4) , 1949, Ibid., 8,1948. 

(5) , 1950, Ibid., 9, 1949. 

(6) , 1951, Ibid., 10, 1950. 

(7) ., 1952, Ibid., 11, 1951. 

(8) , 1953, Ibid., 12, 1952. 

(9) - , 1955, /AW., 14, 1954. 

(10) , 1956, Ibid., 15, 1955. 

(11) Chapman, V.J., 1950, The Marine Algal 

Communities of Stanmore Bay, New 
Zealand. Studies in Intertidal Zonation 1, 
Pac. Sci. 4 (1): 63-68, 3 figs., 1 table. 

(12) Coleman, J., 1933, The Nature of Inter- 

tidal Zonation of Plants and Animals, 
Jour. Mar. Biol. Assoc. U.K., 18: 435-476, 
15 figs., 6 tables. 

(13) Dawson, E.Y., 1945, Marine Algae Asso- 

ciated with Upwelling Along the North- 
western Coast of Baja California, Mexico, 
Bull South. Calif. Acad. Sci., 44(2): 
57-71, pis. 20-22. 

(14) , 1951, A Further Study of 

Upwelling and Associated Vegetation 
along Pacific Baja California, Mexico, 
Jour. Mar. Res., 10 (1): 39-58, 6 figs., 
1 table. 



(15) Doty, M.S., 1946, Critical Tide Factors 

that are Correlated with the Vertical 
Distribution of Marine Algae and Other 
Organisms along the Pacific Coast, 
Ecology, 27: 315-328, 6 figs. 

(16) Ekman,S., 1935,TiergeographiedesMeeres. 

Leipzig, xii + 542 pp., 244 figs. 

(17) Feldmann, J., 1937, Recherches sur la 

Vegetation Marine de la Mediterranee. 
La Cote des Alberes, Rev. Algol., 10: 
1-339, 1 map, 20 pis., 25 figs., 10 tables. 

(18) Gail, F.W., 1918, Some Experiments with 

Fucm to Determine the Factors Control- 
ling Its Vertical Distribution, Publ. 
Puget Sound Biol Sta. 9 2: 139-151, 6 
tables, 1 chart. 

(19) _,19 19, Hydrogen Ion Concentra- 
tion and Other Factors Affecting the 
Distribution of Fucm, Ibid., 2: 287-306, 
pis. 51-52, 4 tables. 

(20) _, 1922, Photosynthesis in Some 

of the Red and Brown Algae as Related 
to Depth and Light, Ibid., 3: 177-193, 
pis. 31-33, 9 tables. 

(21) Hutchinson, A.M., 1949, Marine Plants of 

Economic Importance of the Canadian 
Pacific Coastal Waters, Proc. Seventh 
Pac. Sci. Congr., 5: 62-66. 

(22) Isaac, W.E., 1935, The Distribution and 

Zonation of Marine Algae on the Coasts 
of South Africa, Brit. Assoc. Adv. Set., 
Kept. Ann. Meet. 1935: 455-, London. 

(23) Lamouroux, J.V.F., 1825, Distribution 

G6ographique des Productions Aqua- 
tiques. Hydrophytes des eaux Salees. 
Dictionnaire Classique d'Hist., not. de 
Bory de Saint-Vincent, 7:245-251, Paris. 

(24) _-, 1826, Memoire sur 

la Geographic des Plantes Marines, Ann. 

(25) Okamura, K., 1926, On the Distribution 

of Marine Algae in Japan, Proc. Third 
Pan-Pacific Sci. Congr., 1:958-963, 1 

(26) , 1932, On the Nature of the 

Marine Algae of Japan and the Origin of 
the Japan Sea, Bot. Mag. Tokvo, 41: 

(27) Scagcl, R.F., 1948, An Investigation on 

Marine Plants near Hardy Bay, B.C. 
Prov. Dept. Fish. No. 1, 70 pp., 11 
tables, 26 figs. 

(28) - - , 1957, An Annotated List of 
the Marine Algae of British Columbia 
and Northern Washington, Bull. Nat. 
Mus. Can., 152, 286 pp., 1 fig. 

(29) Setchcll, W.A., 1893, On the Classification 

and Geographical Distribution of the 
Laminariaceae, Trans. Connect. Acad. 
Arts and Sci., 9:333-375. 

(30) .. _- _, 1917, Geographical Distri- 

bution of the Marine Algae, Science, 45 : 

(31) _, 1935, Geographic Elements 

of the Marine Flora of the North Pacific 
Ocean, Amer. Nat., 69: 560-577. 

(32) Stephenson, T.A., and Stephenson, A., 

1949, The Universal Features of Zonation 
between Tide-marks on Rocky Coasts, 
Jour. Ecol., 37 (2): 289-305, pi. 8, 4 figs. 

(33) Tokida, J., 1954, The Marine Algae of 

Southern Saghalien, Me. Fac. Fish, 
Hokkaido Univ., 2 (1): 1-264, 15 pis., 
4 figs., 5 tables. 

(34) Womersley, H.B.S., 1956, The Marine Algae 

of Kangaroo Island, IV: The Algal 
Ecology of American River Inlet, Austra- 
lian Jour. Mar. and Freshwater Res., 
7(1): 64-87, 7 pis., 4 figs. 


G.F. PAPENFUSS: Is anything known concerning growth 
rates of the larger brown algae in the North Pacific? 

F.R. SCAGEL: In Nereoaplis during July and August, 
the blades can increase in length by 2 to 3 inches per day. 

Small sporophytcs a few inches in height in April can de- 
velop into mature, reproducing plants of over 100 feet 
in length by late June of the same year. 





Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, U.S.A. 


What is the maximum possible rate of organic 
production in nature? Is this maximum ever 
attained in the ocean? What are the limiting 
factors of the marine environment which normally 
prevent this potential from being realized or 
even approached? 

These fundamental questions may be attacked 
in either of two ways. First, through extensive 
and repeated oceanographic surveys we may 
accumulate a body of data pertaining to the 
magnitude of primary production in the sea. 
Eventually these data should include the appro- 
ximate range of this process, and through correla- 
tion with chemical and physical characteristics 
of the water, one should be able to extrapolate 
or predict production values for similar oceanic 

Second, we may consider the efficiency of 
utilization of radient energy in the photosyn- 
thetic process and in the organic yeild of mass 
algal cultures, the environmental conditions which 
influence and control this efficiency, and the exis- 
tence and relative importance of these factors in 
the sea. In this way, too, it should be possible 
to predict the potential and, if sufficient informa- 
tion is available, the actual magnitude of primary 
production in the various parts of the ocean as a 
function of its physical and chemical characteris- 

Both of these methods have obvious disad- 
vantages and limitations, so an attempt will be 
made to use a combination of the two in the 
following discussion. First the magnitude and 
efficiency of photosynthesis will be reviewed 
throughout its entire measured range starting 
with carefully controlled quantum yield experi- 
ments in the laboratory, through mass algal cul- 
ture experiments, both indoors and out, to meas- 
urements of primary production in the sea, 
from the highest known values to the lowest. 
This will be followed by a consideration of the 
factors which affect photosynthetic efficiencies, 
treating these quantitatively as far as is possible. 
Finally an attempt will be made to evaluate the 
relative importance of these factors in the various 
parts of the ocean and to relate them to the 
observed values of primary production. 



The quantum requirement or quantum effi- 
ciency of photosynthesis (the reciprocal of the 
quantum yield) may be defined as the number of 
quanta of light energy required to reduce one 
mole of CO 2 to CH 2 O. Although a minority of 
investigators, mainly Warburg and Burk (28), 
have reported extremely low quantum require- 
ments (4 or less approaching 100% efficiency 
of utilization of the absorbed light), it is the opi- 
nion of most plant physiologists that a maximum 
sustained photosyntetic yield requires 8-12 
quanta of red light for the reduction of one 
mole of CO 2 . (See 3, 30, 5, 6, 2.) Values within 
this range have been observed in a wide variety of 
plant species, the literature on which is extensive- 
ly reviewed by Rabinowitch (77, chapter 29) 
and Daniels (2) . 

As a quantum of red light contains 41 k cal 
and one CH 2 O group has a heat of combustion 
of 112 k cal, an average quantum requirement of 
10 corresponds to an efficiency of 

(112x 100)/(lOx41) = 27%. 

Efficiencies of this general order are obtained 
only at very low intensities of light and when the 
physiological condition of the plants and their 
external environment are considered to be optimal 
for photosynthesis. Presumably, then, this (appro- 
ximate) value represents the maximum biological 
potential of plants in converting light into chemi- 
cal energy, a maximum which is imposed by 
the limitations of the physical-biochemical pro- 
cess of photosynthesis. 


Experiments in mass algal culture by the 
Solar Energy Research Group at Agricultural 
College, Wageningen, Netherlands, have demon- 
strated that the maximum sustained efficiency 
observed in photosynthesis experiments is closely 
approached in terms of the organic yield of rela- 
tively long-term growth experiments. (See 
11, 30, 29, 27.) Mass cultures of Chlorella 
were grown by this group under artificial illumi- 
nation with yield efficiencies, calculated on the 



basis of the heat of combustions of the cells 
produced, frequently ranging from 10-15% and 
occasionally greater than 20%. As the organic 
yield in such experiments does not take into 
consideration respiratory losses which are auto- 
matically corrected for in manometric photo- 
synthesis measurements, the proximity of some of 
their efficiency values to that observed in quantum 
yield studies is truly remarkable. 

However, when these cultures were grown out 
of doors under natural illumination, efficiencies 
declined markedly to values of 1-3% under full 
sunlight, as high as 6% in shaded cultures (22% 
in sunlight) 1 . 

Somewhat greater yields were obtained by a 
group of Japanese workers (13) who grew Chlo- 
rella (and other unicellular algae) throughout 
the year in outdoor bubbling cultures. Their 
yields, in grams dry weight of organic matter 
produced per square meter per day, averaged 
16.5 in summer, about 2.4 in winter when 
temperatures were apparently unfavorable. If 
we may assume that their Chlorella were similar 
in composition to the "normal" cells of Ket- 
chum and Redfield (9) its heat of combustion 
calculated by indirect colorimetry would be 5.4 k 
cal/gram, which is comparable to the values 
reported by Kok (11). Using this figure, their 
yield data, and the average radiation values 
for each experiment, a wide range of efficiencies 
were obtained from 0.06% to 14.0%, averaging 
about 4%. To the author's knowledge, the 
maximum yields obtained by this group represent 
the most efficient photosynthetic utilization of 
natural illumination which has been reported. 




We may now turn to the ocean and present a 
brief and partial review of primary production 
values in various coastal and deep sea areas. No 
attempt will be made to review the literature 
completely, and all earlier values of uncertain 
validity are omitted. In general an attempt has 
been made to represent as many different types of 
marine environments and as wide a range of 
production values as possible. Many of the data 
are as yet unpublished. 

Several different methods have been used to 
measure or calculate production by the various 
authors. These will not be described here, but 
the reader may refer to the cited references for 
this infomation. Odum (14), Odum and Odum 
(15), and Riley (19) have based their values on 
measurements of in situ oxygen changes in the 
water, using quite different techniques in each 
case. Yentsch used the familiar "light and dark 
bottle" method of measuring oxygen production 
in his East Sound, Washington, studies. Stee- 
mann Nielsen (25 ) and, in part, the present 
author employed the C 14 method (24). The 
remainder of the values were calculated from 
radiation, light penetration, and chlorophyll data 
using the method recently described by Ryther 
and Yentsch (23). 

Radiation values, except where specifically 
measured with a standard type radiation meter, 
were taken from Kimball (JO) who gives mean 
radiation values at sea level on a world-wide and 
seasonal basis corrected for average cloud cover. 

The use of Kimball's tables many be somewhat 
unsatisfactory in estimating efficiencies of pro- 
duction measured by the C 14 method as the latter 
were obtained on specific days when the radiation 
may have deviated considerably from the mean. 
This objection does not apply to values obtained 
by the chlorophyll-light method which themselves 
were calculated by using Kimball's tables. All 
other efficiencies were based on direct measure- 
ments of radiation except for Odum (14) and 
Odum and Odum (15) who estimated radiation 
from the equation given by Kennedy (8). 

The organic yield (g carbon/m 2 /day) has been 
calculated from oxygen production using an 
assimilatory quotient of 1.25 (see 22) which 
represents organic carbon or approximately 
50% of the ash-free dry weight of plant material. 
This may be converted into comparable energy 
units by multiplying by 2 x 5.4, assuming that 
the heat of combustion of the organic yield is the 
same as that of normal Chlorella cells or 5.4 k 
cal/g (see above). 2 Efficiency was then calculated 
by dividing the daily organic production by 50% 
of the incident radiation, both expressed as gram 
calories/cm 2 /day. 

Table 1 gives the seven highest single values for 
primary production known to the author for the 
marine environment. Of these, the highest values 

1 Photosynthetic efficiencies of the utilization of solar energy are calculated here and elsewhere in this paper on the basis 
of the visible spectrum only (400-700 mu) or roughly 50% of the total incident radiation. 

2 If production is calculated as glucose-carbon (assimilatory quotient = 1), a higher value for carbon assimilation is 
obtained. This may be multiplied by 2.5 (as glucose is 40% carbon) x 3.7 (the heat of combustion of glucose) giving 
nearly the same energy content as of calculated on the basis of the actual composition of the plants. 



were found in the benthic populations studied by 
Odum (14) and Odum and Odum (IS), 10.3 g 
carbon/m 2 /day for the turtle grass community 
and 9.8 g carbon /m 2 /day for the coral reef, cor- 
responding to efficiencies in the utilization of the 
radiation reaching the surface of the water of 
4.0% and 3.1% respectively. 

The other production values in Table 1 range 
from 2.8 to 5.4 g carbon/m 2 /day with less than 
two-fold variation. The implications of the 
striking similarity of these values, obtained 
from widely differing types of environments, will 
be discussed in the final section of this paper. 

In contrast to the high values reported in 
Table 1, Table 2 shows what may be considered 
as average or normal production rates for various 
estuarine, inshore, coastal, and oceanic areas. 
Again the most striking feature of these is their 
similarity. The shallow water areas have pro- 
duction values ranging from about 0.1 to 1.5 g 
carbon/m 2 /day, and it is interesting to note that 
the same range was observed seasonally in the 
two regions where production was followed 
throughout the year, Long Island Sound and the 
continental shelf off New York. 

It is perhaps surprising that the productivity 
of a polluted embayment, Great South Bay, 
averaged only 0.24 g carbon/m 2 /day in midsum- 
mer despite the fact that chlorophyll concentra- 
tions in the euphotic layer averaged about 

10 mg/m 3 or some 10-20 times that of unpolluted 
coastal areas. This is probably due to the 
shallow depth of Great South Bay and the high 
turbidity of its waters, factors which will be dis- 
cussed in more detail later. 

Production measurements in Allen Bay, in the 
Canadian Arctic made by Mr. Spencer Apollonio 
almost daily during the brief open-water season 
(July-August) showed no indication of an initial 
flowering of any magnitude following the breakup 
of the ice, though values did decrease throughout 
the period from 0.69 to 0.02 g carbon/m 2 /day, 
averaging 0.19. One possible explanation for the 
low production of this area is that much of the 
euphotic zone was diluted with melt water low in 
nutrients concentration. 

The oceanic values for the North Atlantic were 
obtained during a passage from Woods Hole, 
Mass, to Plymouth, England by HMS Discovery 
III in April, 1957. They are probably higher 
than average for the year as the cruise apparently 
coincided with the spring diatom flowering (see 
the deep water continental shelf values for April 
in comparison with the rest of the year). The 
four high values from the Grand Banks were 
obtained during the same cruise. 

However, five measurements from the Sargasso 
Sea made by Atlantis during the same period 
showed no evidence of a spring maximum in those 

Table 1. 
Some maximal values and efficiencies of primary production in coastal and oceanic waters. 







g C/m 2 / 



Turtle grass 
Coral reef 

Long Key, Fla. 
Japton Reef, 
Eniwetok Atoll 

Odum (1957) 
Odum & Odum 

Aug., 1955 
July, 1954 


In situ 
In situ 



Spring flowering 

Grand Banks 


Ryther & Yentsch 

Apr., 1957 

380*** Chi 



Polluted estuary 

Forge River, 
Moriches Bay, 
L. I., N. Y. 

Ryther & Yentsch 

Aug., 1956 






East Sound, 


July, 1954 







Walvis Bay, 

Steemann Nielsen 

Dec., 1950 





Spring flowering 

Cont. Shelf 
off N. Y. 

Ryther & Yentsch 

Apr., 1957 






* From Kennedy (1949); ** measured; *** From Kimball (1928). 


Table 2. 

Magnitude and efficiency of primary production in selected coastal and oceanic waters. 

_ j _ 






Reference Date 

(g cal/ 


Prod, (g 
m 2 /day) 



cm 2 /day) 




Ryther &Yentsch 

Aug., 1956 








Bay, L. I., 



Long Is. 


Riley (1956) 



In situ 1.06 



Sd., N.Y. 

May-Aug., 1952 




Aug.-Nov., 1952 


i 1.22 


jNov.-Feb., 1952-3 




Feb.-Mar., 1953 





Allen Bay, 



July- Aug., 1956 







Ryther & 

Is., N.W.T. 



Cont. Shelf 

Off N.Y. 


Ryther & 

Sep., 1956 





(25 m depth) 


Dec., 1956 





Feb., 1957 




Mar., 1957 




Apr., 1957 




Cont. Shelf 

Off N.Y. 


Ryther & Yentsch 

Sep., 1956 





(500 m depth) 


Dec., 1956 




Feb., 1957 




Mar., 1957 




Apr., 1957 




Cont. Shelf 



Ryther & Yentsch 

Apr., 1957 






S. Atlantic 


Steemann Nielsen 

Dec., 1950 

476 O4 






i i 




Ryther & Apr., 1957 

380 Chi 1 0.47 





(unpub.) : 



Turner, Ryther 

Apr., 1957 

477 Chi 




& Yentsch 





Steemann Nielsen 

June, 1952 










Ryther (unpub.) 

Feb., 1955 


Ci4 0.43 






Steemann Nielsen 

Mar., 1952 


O4 0.15 








C" | 0.14 













> *> 

| 410 









waters, the mean of 0.044 g carbon/m 2 /day 
agreeing almost exactly with Steemann Nielsen's 
values for the Sargasso Sea (0.048) obtained by 
the C 14 value during June, 1952. Actually these 
values varied by almost ten-fold from a minimum 
of 0.009 to a maximum of 0.081. 

The remaining oceanic values, from Ryther and 
Steemann Nielsen, were obtained by the C 14 
method and hence are probably somewhat lower 
than would have been given by other methods. 
In spite of this, these data indicate (as do the 
majority of Steemann Nielsen's values not 
included here) that oceanic production is consider- 
ably lower than that of coastal and inshore 
waters. However, this cannot be considered as a 
final conclusion until substantiated by one or 
more studies of the annual cycle of primary 
production in a truly oceanic area. 

Efficiencies of primary production of inshore 
waters range from about 0.1 to 1.0 per cent, 
again showing this range seasonally as well as 
regionally. Efficiencies for oceanic production 
are less certain. They appear to be lower, but 
also vary by at least an order of magnitude, 
ranging from 0.02 to 0.27% in the few data 
presented here. 

We may now summarize the rough averages of 
photosynthetic efficiencies from all sources dis- 
cussed above, from quantum-yield laboratory 
experiments to the Sargasso Sea. Listed in 
decreasing order of magnitude, they appear as 

Quantum yield of photosynthesis 27.0 
Indoor algal culture yields 15.0 

Outdoor algal culture yields 3.0 

Maximal marine values 2.2 

Average coastal and inshore 

waters 0.5 

Average oceanic waters 0. 1 

In the following discussion an attempt will be 
made to review and evaluate the environmental 
factors responsible for the decline of efficiency 
from the top to the bottom of this list. 



In quantum efficiency experiments, photosyn- 
thesis is corrected for the effects of respiration. 
However, in the mass algal culture work described 
above, the yield of organic matter represents the 
net effect of both photosynthesis and respiration. 
Similarly the C 14 method for estimating primary 
production in the sea appears to measure photo- 


synthesis minus respiration (see 20). The C 14 
values given in Tables 1 and 2 which were ob- 
tained by the author have not been corrected 
for respiration while those of Steemann Nielsen 
(25) have been increased by 4% to allow for res- 
piratory loss. The other values reported above 
were obtained by methods which correct for res- 
piration, and this section does not apply to them. 

Respiration is probably never less than 5-10% 
of maximum sustained photosynthesis with light 
and all other factors optimal. In an earlier 
paper, the author showed the relationship be- 
tween total daily photosynthesis within the 
entire euphotic layer and the incident radiation 
falling on the surface of the water (21). This 
curve is reproduced in Fig. 1. If one assumes 
that respiration is 7.5% of optimal photosyn- 
thesis (taking the middle of the range suggested 
above), the curve for total daily respiration 
within the euphotic layer relative to photosynthe- 
sis may be shown as a straight line (i.e., indepen- 
dent of radiation), as in Fig. 1 . The ratio of res- 
piration to photosynthesis, given as a percentage, 
then represents the respiratory loss. 

It may be seen that with an incident total 
radiation of 100 g cal/cm 2 /day or less the plants 
are at or below compensation. With the highest 
radiation values reaching the surface of the earth, 
respiratory losses are barely less than 30%, and 
within the range normally encountered over most 
of the earh (250-500 g cal/cm 2 /day), the respira- 
tory loss lies between about 30% and 50%. 

These losses apply only to plants within the 
euphotic layer (the depth of penetration of 
approximately 1 % of full sunlight). If part of 
the plant population under consideration (wheth- 
er an algal culture, a phytoplankton population, 
or a bed or attached algae of rooted plants) 
extends below this depth, these plants will impose 
an additional respiratory loss to the system. 
Thus in an unstable water column with a phyto- 
plankton population which is wind-mixed to twice 
the depth of the euphotic zone, net growth of the 
population cannot occur if the incident radiation 
is less than about 275 g cal/cm 2 /day. 

Similarly a dense culture of Chlorella, although 
brightly illuminated at its surface and supplied 
with an excess of nutrients and CO 2 , may cease to 
grow not because photosynthesis is stopped, but 
because the light cannot penetrate to sufficient 
depth to permit photosynthesis to compensate 
for the respiratory loss of the culture as a whole. 

The relationship between photosynthesis and 
respiration shown in Fig. 1 is based upon natural 
daylight conditions and therefore does not apply 



200 JOO 400 500 600 TOO 

TOTAi INCIDENT RADIATION (fl col / em f doy ) 

Fig. 1 . Total daily photosynthesis (P), respiration (R), and 
the ratio of respiration to photosynthesis ( % respiratory 
loss) of phytoplankton within the euphotic zone as a func- 
tion of total daily incident radiation. 

to cultures constantly illuminated by artificial 
lights. Nevertheless it is clear that a large por- 
tion of the cells of these cultures are always in 
sub-optimal light if not in darkness, and it seems 
reasonable to assume that the discrepancy 
between the efficiency of yields of algal cultures 
grown under artificial illumination (15%) and the 
quantum efficiency of photosynthesis (27%) may 
be due to this factor alone. 

Similarly the C 14 values reported by the 
author are only 1/2 to 3/4 as high as the true 
values for total photosynthesis, and Steemann 
Nielsen's values including a 4% correction for 
respiration are probably also low. 


In quantum yield experiments, care is taken to 
consider the utilization of only that light which 
is actually absorbed by the plants. This cannot be 
easily determined in culture yield experiments or 
in measurements of natural photosynthetic 
rates, and it has sufficed for most workers to base 
efficiencies on the utilization of the light falling 
on a square unit of water surface. This introduces 
a rather small but significant error in that a frac- 
tion of the incident radiation is lost through 
reflection from the surface or from back scatter- 
ing out of the water. 

Powell and Clarke (16) measured these losses 
from the sea surface. At solar angles greater 
than 30 they found that about 4% of the incident 
light was lost through the combined effects of 
reflection and back scattering on clear days, 
about 6.5% on cloudy days. This appears to be 

independent of the sea state, from flat calm to 
winds strong enough to produce white caps. 

Reflection losses increase greatly at solar angles 
below 30, as predicted by theory, but as relatively 
little absolute radiation reaches the earth's surface 
at such low solar altitudes, these higher reflection 
values may be discounted. It would appear, 
therefore, that 5% is a reasonable average value 
to assign to this loss. 

Fig. 2. 

A) The quantum requirement of photosynthesis as a func- 
tion of wave length for Chlorella (after Emerson and 
Lewis, 1943) and for Nitzschia (after Tanada 1951). 

B) Energy per mole Quantum of light as a function of 
wave length. 

C) The efficiency of photosynthesis as a function of wave 




It was pointed out that sustained photosyn- 
thesis under carefully controlled laboratory 
conditions requires 8-12 quanta of red light to 
reduce one mole of CO 2 , corresponding to a 
mean efficiency of 27%. These experiments 
are usually carried out at or near a wave length of 
680 mji, the maximum absorption peak for 

Emerson and Lewis (4) and Tanada (26) have 
studied the quantum requirement of photosyn- 
thesis at different wave lengths using Chlorella 
and Nitzschia respectively. Their curves are 
reproduced in Fig. 2A. The differences between 
the two species may not be significant as tech- 
niques may have differed somewhat, but in each 
case the minimum requirement falls within the 
range given above, remains relatively constant 
between about 680 and 550 m^i, increases to a 
maximum at 490 mu, and then partially recovers 
between 490 and 400 m|i. Above 685 mjj, the 
quantum requirement increases markedly for 
both species. 

Figure 2B shows the energy per quantum of 
light between 400 and 700 mji illustrating the 
fact that this drops from a maximum of 71 k cal 
per mole of quanta at the blue end to a minimum 
of 41 k cal at the red end of the visible spectrum. 

As red light is most efficient in terms of its 
energy content per quantum and as the quantum 
requirement of photosynthesis increases in blue 
light, the efficiency of photosynthesis decreases 
from its maximum (25-30%) in red light to a 
minimum of 12.5% in blue light, as shown in 
Fig. 2C. 

The spectral distribution of daylight varies with 
solar altitude and with the water vapor, car- 
bon dioxide, and dust content of the atmos- 
phere. Fig. 3 shows the spectral distribution of 
daylight under average atmospheric conditions 
with an air mass of 2 (solar angle=60) as given 
by Moon (12). If the curves for the two organ- 
isms shown in Fig. 2C are averaged and the mean 
efficiency calculated for the entire visible spec- 
trum, weighted for the spectral distribution of 
sunlight as given in Fig. 3, this value turns out 
to be 18.4%. Thus average solar radiation is 
only about 68 % as efficient for photosynthesis as 
red light of 680 mu. 

In extremely turbid waters, particularly those 
containing dissolved pigmented organic matter 
(the so-called "yellow substance" of Kalle, 7) 
blue and green light may be selectively absorbed 
which would result in somewhat higher efficiencies 


Fig. 3. The spectral distribution of daylight under average 
atmospheric conditions with air mass-2 (solar angle=60). 

6 8 10 


Fig. 4. 

A) Photosynthesis of marine phytoplankton as a func- 
tion of light intensity (after Ryther, 1956a). Broken line 
is the extrapolation of the linear portion of the solid 
line, representing hypothetical sustained maximum pho- 
tosynthetic efficiency. 

B) Efficiency of photosynthesis as a function of light 
intensity, as calculated from Figure 4 A, assuming a max- 
imum efficiency of 100%. 



in the utilization of the light which penetrates to 
greater depths. However, in normal, clear 
coastal and oceanic waters, red light is selectively 
absorbed and scattered by the water, and the 
blue and green penetrate to the greatest depths 
where they would be used still less efficiently than 
the incident daylight discussed above. 


The exact mathematical relationship between 
light intensity and photosynthesis is rather contro- 
versial (see 27), but an examination of many 
intensity-photosynthesis curves (as in 77, chapter 
28) leaves little doubt that the relationship closely 
approximates linearity up to the saturation inten- 
sity. Above this, photosynthesis does not 
increase further, and at intensities 1/4 to 1/3 of 
full sunlight and above it may become severely 
depressed (see 21). 

As soon as the intensity-photosynthesis curve 
departs from linearity, efficiency begins to 
decrease. This may occur at intensities as low 
as 1/20 of full sunlight. 

The laboratory experiments described above 
giving an 8-12 quantum requirement were all 
carried out at low intensities on the linear portion 
of the curve. If this linear portion is extrapo- 
lated, one may then calculate efficiencies at any 
intensity by comparing the extrapolated line with 
the actual photosynthesis curve. This has been 
done in Fig. 4A using the photosynthesis-intensity 
curve for marine phytoplankton given by Ryther 
(21). The broken line obtained by extrapolating 
the linear portion of the curve, represents the 
hypothetical sustained maximum rate of photo- 
synthesis. The efficiency, obtained from the 

00 300 400 500 00 


Fig. 5. Daily efficiency of photosynthesis by phytoplank- 
ton within the euphotic zone as a function of total daily 
radiation, assuming a maximum efficiency of 100%. 

Fig. 6. The fraction of light absorbed by water as a func- 
tion of the depth of the euphotic zone, defined as the 
depth of penetration of 1 % of maximum sunlight incident 
to the surface. 

ratio of corresponding points on the two curves, 
is shown in Fig. 4B, with the maximum efficiency 
arbitrarily given a value of 100%. 

From Fig. 4B and instantaneous values for 
total incident radiation (corrected for reflection 
loss), one may calculate photosynthetic efficien- 
cies throughout the day, and these may be aver- 
aged to give the mean daily efficiency. This, 
however, will characterize only those organisms 
at the surface of the water. The deeper, shaded 
plants will operate at higher efficiencies. But as 
light is absorbed exponentially, the greater part 
of the incident radiation is absorbed at the 
higher, less efficient intensities. To evaluate this 
quantitatively it is necessary to calculate mean 
daily efficiencies at several depths where the surface 
intensity is decreased by known amounts, weight 
each depth interval according to the amount of 
radiation absorbed within that interval, and 
integrate over the entire illuminated water 

This was done for eight days, each character- 
ized by a different incident radiation ranging from 
about 30 to 800 g cal/cm 2 /day. The resulting 
curve showing the relationship between total 
incident radiation and daily photosynthetic 
efficiency for the antire phytoplankton popula- 
tion is given in Fig. 5, assuming a maximum 
efficiency of 100%. 

According to these calculations, photosynthe- 
tic efficiencies begin to decrease when the incident 
radiation exceeds about 50 g cal/cm 2 /day, and 
falls to 40% of its maximum on days of highest 
radiation. Under normal conditions in tropical 
and temperate regions, where average radiation 



values fall between 250 and 500 g cal/cm 2 /day, 
efficiencies are reduced to 70-50% of their 
biological potential from the effects of light 
intensity. Although these calculations are based 
upon the photosynthesis-light intensity relation- 
ships of marine phytoplankton, they probably 
apply reasonably well to marine plants in general. 


The availability of plant nutrients has a pro- 
nounced effect upon the rate and efficiency of 
photosynthesis, a fact with which plant physiolo- 
gists and ecologists are equally well acquainted. 
A single example is perhaps sufficient to illustrate 
this point. In an earlier paper (20), an experi- 
ment was described in which the marine flagellate, 
Dunaliella euchlora, was grown in a culture 
medium with a limiting supply of nitrogen and 
phosphorus, the two elements which are probably 
most frequently limiting to photosynthesis in the 
sea. The results of this experiment are repro- 
duced in Table 3. Exponential growth of the 
flagellate occurred for the first five days after 
which essentially the same cell concentration 
persisted for the next twenty three days. Photo- 
synthesis per cell decreased twenty-fold from its 
maximum on the second day to its minimum on 
the twenty-eighth day. Respiration decreased to 
one-half its maximum during the same period. 
The two processes were equal and the culture at 
compensation level by the end of the experiments. 

In nutrient deficient plants, particularly those 
limited by nitrogen, the chlorophyll content of the 
cells decreases markedly. If these cells absorbed 
less light and the latter could thereby penetrate 
to greater depths, photosynthetic efficiency per 

Table 3. 

Photosynthesis (P) and respiration (R) 

in a nutrient-deficient pure culture of Dunaliella 

euchlora expressed in ml O 2 /24 hrs. 

(From Ryther, 1954.) 


Mean cell count 
(10^ cells/liter) 



1- 2 


2- 3 


3- 4 


4- 5 


5- 6 


6- 7 




28-29 | 415 










(xlO 9 ) 








unit of surface area would not be reduced to 
the same extent as when calculated per unit 
volume. The following section will have some 
bearing on this question. Actually, however, 
this is not the case because the concentration of 
the carotenoid pigments decreases very slowly 
in starved cells and their light absorptive charac- 
teristics do not differ greatly from that of healthy 
plants. This was determined by measuring the 
light absorption of a suspension of Dunaliella 
in the 10 cm light path of a Beckman DU 
Spectrophotometer, averaging the percent trans- 
mission at all wave lengths between 400 and 700 
n\[i. This varied from 84% in a healthy, growing 
culture to 88% in a nitrogen-starved culture 
which had not grown for fourteen days. 

Thus under the conditions of this experiment, 
the efficiency of photosynthesis decreased to 5 % 
of its maximum from the effects of nutrient defi- 
ciency. If photosynthesis were measured by a 
method which does not correct for respiration, 
this effect would be still more striking as the 
organisms at the end of the experiment, which 
were at compensation level, would indicate no 
photosynthesis whatever. 

While it is perhaps doubtful that plants in this 
extremely impoverished condition normally exist 
in the ocean, nutrient deficiency is still probably 
the most important single factor limiting the 
efficiency of photosynthesis over most of the 
marine environment as will be discussed later. 




Plants living in an aqueous medium are at 
some disadvantage over terrestrial forms as some 
of the incident radiation must be absorbed by the 
water itself. According to Jerlov, the extinction 
coefficient of pure sea water is .033. This loss is 
negligible if the plants are located near the surface 
and sufficiently concentrated so that the light is 
all absorbed within a few meters. In a sparse 
plankton population, however, as is typical of the 
open sea, light may penetrate to much greater 
depths and a considerable fraction of it is then 
absorbed by the water rather than the plants. 
Fig. 6 shows the percentage of the incident light 
which is absorbed by the water as a function of 
its transparency. From this curve it is obvious 
that in such a place as the Sargasso Sea, where the 
euphotic zone may exceed 100 meters, over 80% 
of the light is absorbed by the water, and the 
efficiency of the photosynthetic utilization of the 


radiation incident to the sea surface is corres- 
pondingly lowered. 

This, however, cannot be considered as a direct 
cause of decreased photosynthetic efficiency, 
as a sparse plankton population itself merely 
reflects the decrease or absence of photosynthesis 
and plant growth which in turn must ultimately 
result from a deficiency of plant nutrients or 
light. It is mentioned here to point out the fallacy 
in the concept that production per unit of surface 
area should be comparable in dense and sparse 
plankton populations, an idea based on the 
assumption that the lower rate of photosynthesis 
per unit volume in the latter case may be com- 
pensated for by a deeper euphotic zone. This 
obviously does not take into consideration ab- 
sorption by the water as Steemann Nielsen (25) 
has pointed out. 

However, there is some justification for this 
concept because in coastal or inshore waters with 
a shallow euphotic layer, a large fraction of the 
light may be absorbed by non-planktonic parti- 
culate matter, either organic or inorganic in 
nature and probably often associated with the 
bottoms. Thus Riley, (19) taking the relationship 
between the extinction coefficient and the chloro- 
phyll content of offshore waters and plankton 
blooms where light absorption by non-planktonic 
particulate matter was assumed to be minimal, 
calculated that an average of only 1/3 of the light 
incident to Long Island Sound is absorbed by 
plants, the remainder being attributed to absorp- 
tion by other particulate matter. 

Additional light may be absorbed in coastal 
waters by dissolved pigmented material (i.e., 
the "yellow substance" of Kalle, 7) which prob- 
ably consists of organic compounds of terrige- 
nous origin and are of relatively little significance 
on the open sea. Finally in extremely shallow 
areas, such as Great South Bay with a mean depth 
of about 2 meters, light may penetrate to and be 
absorbed by the bottom. 

These various factors probably explain why 
production and efficiency values are so similar 
in Great South Bay, Long Island Sound, and the 
continental shelf area, regions where light absorp- 
tion by the water, non-planktonic particulate and 
dissolved matter, and the bottom, although of 
different relative importance in each case, may 
add up to nearly the same total effect. 

While light absorption by water was mentioned 
above as an indirect cause of lowered efficiency, 
symptomatic of other deficiencies, such is not the 
case in the absorption of light by particulate and 
dissolved matter which are quite independent 


of and in competition with the plants for the 
available light. 


This discussion is based upon the premise, 
described in the first section of this paper, 
that the maximum sustained efficiency of photo- 
synthesis is represented by the conversion of 
about 27% of the light absorbed by the plants, 
this light being at or near 680 mji and at an 
intensity well below the saturation level. Turning 
from these ideal laboratory conditions to those 
existing in nature, and considering the utilization 
of sunlight falling on the sea surface by the 
underlying plants, we have seen that wave lengths 
and intensities are no longer optimal, that light is 
lost by reflection and back scattering, by absorp- 
tion by the water itself, by dissolved or particu- 
late matter other than plants, or by the bottom. 
Some of these factors are relatively constant, 
others may vary greatly, but all combine to lower 
efficiencies of natural photosynthesis to a small 
fraction of the "normal quantum yield" as it is 
known to the plant physiologist. 

Let us consider, for instance, the maximum 
possible efficiency in nature by marine plants. 
Under the best of conditions, there is still a 5% 
loss of light by reflection and back scattering, a 
wave length effect resulting in efficiencies some 
32% below that observed in red light, and an 
intensity effect, depending upon the length and 
brightness of the day, reducing efficiencies 30- 
50% below that possible in sub-saturation inten- 
sities. The accumulated effect of these factors is 
to decrease efficiencies 67-87% of the maximum 
potential of 27% or to a level ranging from 
3.5% to 9.0%. 

Are these low values reasonable? They may 
be checked by re-examining the organic yields of 
carefully conducted outdoor mass algal culture 
experiments. Here efficiencies were found to 
average about 2% by the Netherlands group, 
about 4% by the Japaness investigators. These 
yields, however, were not corrected for respira- 
tory losses which, we find, could hardly be less 
than 30% and may reach 100% in extremely 
dense cultures. If we take 50% as an average 
respiratory loss, the efficiency of photosynthesis 
in these mass cultures would be 4-6%, or roughly 
the same as that calculated above. 

Are these efficiencies encountered in the sea? 
The data presented in Table 1 shows that they may 
be attained, or at least closely approached in a 
wide variety of different marine environments. 



These, however, all have one feature in common, 
a sufficiency of nutrients. The flowing-water, 
benthic population appear to live under the most 
optimal conditions, and the reasons for this are 
obvious -they live at the bottom of a shallow, 
clear column of water which removes very little 
of the light, and they are bathed in a constantly 
replenished medium from which the nutrients 
can never become exhausted. 

Similarly polluted regions are characterized by 
the constant addition of nutrients from an 
external source. Here, however, the existence of 
dissolved or particulate substances competing 
with the plants for light is almost a certainty, 
as pollution by definition is of terrestrial origin 
and highly productive polluted areas are virtually 
confined to harbors, estuaries, etc., which nor- 
mally contain considerable amounts of such 
substances. Furthermore, most sources of 
pollution also contribute substantial quantities of 
light absorbing material (i.e., organic detritus, 
bacteria, silt, dyes, etc.) to the water in addition 
to the essential plant nutrients. Therefore it is 
unlikely that such an environment would sustain 
production at efficiencies as high as those possible 
in the flowing-water benthic communities. 

More favorable conditions for photosynthesis 
may exist in coastal and offshore areas where the 
nutrients originate through upwelling or mixing 
processes from the rich, deep ocean waters which 
are relatively free of light absorbing substances. 
However, high production rates under such 
circumstances arc, for the most part, brief and 
transitory, either in time or space. Although high 
production occurs at or near an area of upwelling, 
it quickly diminishes as the upwelled water 
spreads over the ocean surface. Production may 
remain high at a given geographical area, but it 
is not sustained in a given volume of water or by 
a given population of plants. Similarly, most 
plankton blooms or flowerings are temporary, 
usually representing the culmination of mixing 
or regeneration processes during a period when 
plant growth can not occur (e.g., during temperate 
or polar winters). With the return of favorable 
conditions for photosynthesis, the temporarily 
enriched waters may support a brief and dramatic 
period of extremely high production. 

According to Red field ( 18), the relative propor- 
tion of carbon, nitrogen, and phosphorus in 
plankton is 100:15:1 by atoms. A production 
rate of 5 grams of carbon per day within a 10 
meter water column (such high production 
rates result in and are therefore restricted to an 
extremely shallow euphotic zone) would therefore 


require 6 ugA of nitrogen and 0.4 ugA of phos- 
phorus per liter per day. There is no deep water 
in the ocean rich enough to meet this demand 
for more than a few days. 

Thus efficiencies of 3-9%, representing the 
maximum photosynthetic potential under natural 
illumination, are rarely encountered in the ocean. 
The average efficiencies reported in Table 2 for 
coastal and offshore waters are, for the most 
part, one to two orders of magnitude below this. 

Two variable environmental factors have been 
discussed, both of which decrease efficiencies 
below this maximum potential. These are (1) 
light absorption by dissolved and particulate 
matter other than plants, and (2) nutrient deficien- 
cies. The former undoubtedly plays an impor- 
tant part in reducing efficiencies in coastal waters. 
As mentioned earlier, Riley (19) has estimated 
that only one-third of the light entering Long 
Island Sound is absorbed by plants. Similar 
calculations by the author have indicated that 
anywhere from 27% to 66% of the light pene- 
trating the waters of the continental shelf is uti- 
lized by the phytoplankton, the remainder being 
about equally divided between other particulate 
matter and the water itself. 

It seems doubtful, however, that this factor 
can be important in the clear waters of the open 
sea, and the conclusion seems inevitable that the 
greater part of the oceans is normally nutrients 
deficient. Unfortunately relatively little is known 
about the concentrations of all the essential 
plant nutrients in the surface water of the ocean; 
their rates of uptake by the phytoplankton, 
limiting concentrations for photosynthesis, and 
rates of regeneration by mixing and decomposi- 
tion. Consequently the relative importance of this 
factor cannot be quantitatively assessed. How- 
ever, the conclusion that nutrient deficiencies are 
responsible for the low efficiencies (compared to 
the potential efficiency of 3% or more) normally 
found in the sea is based upon the following bits 
of indirect evidence or reasoning: (1) such 
nutrient concentrations as have been measured 
in the surface waters of the open sea, mainly N 
and P compounds, are normally extremely low, 
often undetectable. (2) Sustained maximum 
efficiencies such as are possible under natural 
illumination would require the constant replen- 
ishment of nutrients whose only source is the 
ocean depths. This water cannot be mixed 
upward into the surface layers without, at the 
same time, carrying the plants down out of the 
euphotic zone. In most of the oceans, thermal 
stratification is an effective barrier against such 

mixing. (3) The only marine areas where this 
maximum potential efficiency is approached, 
temporarily or permanently, are those situations 
in which some mechanism exists for the enrich- 
ment or continued replenishment of nutrients. 
(4) No other environmental factor appears to 
exist which could decrease photosynthetic efficien- 
cies to a comparable degree. 


(1) Clarke, G. L., and James, H.R., 1939, Labo- 

ratory Analysis of the Selective Absorp- 
tion of Light by Sea Water, J. Optical 
Soc.Amer., 29: 43-55. 

(2) Daniels, F., 1956, Energy Efficiency in Pho- 

tosynthesis, Chapt. 4 in Hollaender, 
A., Radiation Biology, Volume III: 
Visible and Near Visible Light, McGraw- 
Hill, New York. 

(3) Emerson, R., and Lewis, C.M., 1941, Carbon 

dioxide Exchange and the Measurement 
of the Quantum Yield of Photosynthesis, 
Am.J. Bot., 28:789-804. 

(4) _ _. , 1943, The 

Dependence of the Quantum Yield of 
Chlorella Photosynthesis on Wave 
Length of Light, Ibid., 30: 165-178. 

(5) Gaffron, H., 1954, Mechanism of Photosyn- 

thesis, in Autotrophic Microorganisms, 
4th Symp. Soc. Gen. Microbiol., The 
University Press, Cambridge. 

(6) Hill, R., and Whittingham, C. P., 1955, Pho- 

tosynthesis, Methuen and Co., London. 

(7) Kalle, K., 1938, Zum Probleme der Meeres- 

wasserforbe, Ann, Hydrog. u. mar. 
Meteor., 66. 

(8) Kennedy, R. E., 1949, Computation of 

Daily Insolation Energy, Am. Meteor. 
Soc. Bull, 30: 208-213. 

(9) Ketchum, B.H., and Redfield, A. C., 1949, 

Some Physical and Chemical Character- 
istics of Algae Grown in Mass Culture, 
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(JO) Kimball, H. H., 1928, Amount of Solar 
Radiation that Reaches the Surface of 
the Earth on the Land and on the Sea, 
and Methods by which It is Measured, 
Mon. Weath. Rev. Wash., 56: 393-398. 

(11) Kok, B., 1952, On the Efficiency ofChhrella 
Growth, Acta Botanica Neerl, 1: 445- 


(12) Moon, P., 1940, Proposed Standard Solar- 

Radiation Curves for Engineering Use, 
J. Franklin lnst. 9 230: 583-617. 

(13) Morimura, Y., Nikei, T., and Sasa, T., 1955, 

Outdoor Bubbling Culture of Some 
Unicellular Algae, /. Gen. Appl. Micro- 
Mo/., 1: 173-189. 

(14) Odum, H. T., 1957, Primary Production 

Measurements in Eleven Florida Springs 
and a Marine Turtle-Grass Community, 
Limnol. and Oceanogr., 2: 85-97. 

(15) Odum, H.T., and Odum, E.P., 1955, Tro- 

phic Structure and Productivity of a 
Windward Coral Reef Community on 
Eniwetok Atoll, Ecol. Monogr., 25: 291- 

(16) Powell, W.M., and Clarke, G.L., 1936, The 

Reflection and Absorption of Daylight 
at the Surface of the Ocean, /. Optical 
Soc. Am., 26: 1-23. 

(17) Rabinowitch, E. L, 1951, Photosynthesis 

and Related Processes, Interscience 
Publ., II, Part 1, New York. 

(18) Redfield, A. C., 1934, On the Proportions of 

Organic Derivatives in Sea water and 
Their Relation to the Composition of 
Plankton, James Johns tone Memorial 
Vol., Univ. of Liver pool, 176-192. 

(19) Riley, G. A., 1956, Oceanography of Long 

Island Sound, 1952-1954, II: Physical 
Oceanography, Bull. Bing. Oceanogr. 
Coll., 15: 15-46. 

(20) Ryther,J.H., 1954, The Ratio of Photosyn- 

thesis to Respiration in Marine Plankton 
Algae and Its Effect upon the Measure- 
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(21) , 1956a, Photosynthesis in the 

Ocean as a Function of Light Intensity, 
Limnol. and Oceanogr., 1:61-70. 

(22) - - , 1956b, The Measurement of 

Primary Production, Ibid., 72-84. 

(23) Ryther, J.H., and Yentsch, C.S., 1957, The 

Estimation of Phytoplankton Produc- 
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Light Data, Limnol. and Oceanogr., 2. 

(24) Steemann Nielsen, E., 1952, The Use of 

Radio-active Carbon (C 14 ) for Measur- 
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Cons. Internat. Explor. Mer. 9 18: 1 17-140. 

(25) _ _ - , 1 954, On Organic Pro- 

duction in the Oceans, Ibid., 19: 309-328. 



(26) Tanada, T., 1951, The Photosynthetic Effi- 

ciency of Carotenoid Pigments in Navi- 
cula minima, Am. J. Bot., 39: 276-283. 

(27 ) Van Oorschot, J.L.P., 1955, Conversion of 

Light Energy in Algal Culture, Mededel- 
ingen van de Landbouwhogerschool te 
Waginingen, Nederland, 55: 225-276. 

(28) Warburg, O., and Burk, D., 1950, The 

Maximum Efficiency of Photosynthesis, 
Arch. Biochem., 25: 410-443. 

(29) Wassink, E. C, 1954, Problems in the Mass 

Cultivation of Photo-autotrophic Micro- 
organisms, in, Autotrophic micro- 
organisms, 4th Symp. Soc. Gen. Micro- 
bioi, The University Press, Cambridge. 

(30) Wassink, E.C., Kok, B., and Van Oorschot, 
J.L.P., 1953, The Efficiency of Light 
Energy Conversion in Chlorella Cultures 
as Compared with Higher Plants, in, 
Burlew, J. S. : Algal Culture, from 
Laboratory to Pilot Plant, Carnegie Inst. 
Wash., PubL, 600. 







Department of Botany, University of Illinois, Urbana, Illinois, U.S.A. 

Codiwn (order Codiales or Siphonales of the 
green algae) is a relatively large genus (that is, 
compared with other genera of marine algae), 
comprising about eighty species. It is exceed- 
ingly widely distributed, growing on all tropical 
and temperate shores with the notable exception 
of the Atlantic coast of North America north of 
Beaufort, North Carolina 1 . With the cooperation 
of scores of workers throughout the world, 
during the past eight years, I have been able to 
assemble about 2,000 fixed and dried collections. 
In attempting to work out a taxonomic treatment 
of Codium, I began to perceive certain geographic- 
al distributional patterns. It then became im- 
portant to determine whether a standard of 
reasonableness could be established against which 
to judge geographical distributions as a test 
of taxonomic soundness. It was of interest to 
see whether there was justification for being 
suspicious about the alleged occurrence of the 
same species, for example, in the Irish Sea and 
on the California coast. The first step was to 
become aware of previously described distribu- 
tional patterns for seaweeds; the second step was 
to compare these patterns with those that are 
exhibited by Codiwn. 

The field of phytogeography is vast, the litera- 
ture inexhaustible, its data capable of a variety 
of interpretation, extraordinary manipulation, 
and distortion. It has the fascination of a chess 
game. It is a valid, though treacherous, field of 
investigation. While making no claim to com- 
petency as a phytogeographer, 1 believe that 
information divulged by my studies of Codium 
warrants evaluation in general phytogeographic 


It has long been recognized that along most 
coasts temperature is probably the most important 
factor determining the latitudinal distribution 
of marine algae a view particularly expressed 
by Setchell (2, 3, 4), who believed that the 
effect of temperature on reproductive processes 

was critical. Ideally, ocean surface temperatures 
could be expected to vary linearly from the 
equator toward the poles; however, it is well 
known that the major oceanic currents profound- 
ly alter this gradient. Zones of algal distribu- 
tion thus do not coincide with latitudinal zones. 

Figure 1 is a schematic representation (modi- 
fied Mercator projection) of an idealized current 
system, wherein it is assumed that both sides of 
the ocean are delimited by continuous land 
masses bearing due north/south. The pattern of 
currents coincides in many regions with the 
pattern of prevailing winds, but it depends not 
only upon the prevailing winds but upon the 
rotation of the earth and density differences in 
the surface layers. The pattern in the northern 
hemisphere has a mirror image in the southern 
hemisphere. The following features may be 
noted: northern and southern equatorial currents, 
equatorial countercurrent, northern and southern 
high-latitude currents, northern and southern 
high-latitude minor gyrals. Whether a current as 
it bathes a particular shore should be designated 
"warm" or "cold" depends upon the temperature 
that might be expected in the absence of a 
current system. Under the idealized conditions 
of Fig. 1, the western shores of the ocean would 
be warmed by the equatorial currents while the 
eastern shores would be cooled by the returning 
high-latitude currents. Actually, this idealized 
pattern is significantly altered by at least three 
factors: (1) the configuration of major land 
masses, affecting the major current system; 
(2) the configuration of minor land masses, affect- 
ing local gyrals; and (3) upwelling, causing local 
temperature disturbances. 

Figure 2 shows the patterns of currents in the 
Pacific. Because of the funnel formed between 
Asia and North America, the North Equatorial 
Current warms the Japanese coast (as the Kuro- 
shio Current) and its high-latitude extension 
retains enough warmth, despite the admixture 
of cold subarctic water south of the Aleutian 
Islands, to be regarded as a warm current when it 

1 Bouck and Morgan (J) report the discovery at East Marion, Long Island, New York, of Codium fragile &$$. tomentosoides 
(van Goor) Silva, a weed presumably introduced from Europe, where it is becoming increasingly widespread. 



'' ' 


'/>'' L'^??^ 1 * repre f ntati / on of idea . lized current s V stem (modified Mercator projection), a, a' t equatorial currents; 
, b , high-latitude currents ; c, c', equatorial countercurrent ; d t </', high-latitude minor gyrals 



reaches the coast of North America. The north- 
ward moving minor branch, the Alaska Current, 
is definitely warming; the southward moving 
major branch, the California Current, largely 
because of the upwelling of cold water soon 
reaches a point where it no longer has the effect 
of warming, but of cooling. To the northeast 
of Japan, temperatures are depressed by the 
western part of the Bering Sea gyral (the Oyashio 
Current). In the southern hemisphere, the cir- 
cumpolar current of the antarctic region sets the 
stage for a very different pattern. The continent 
of South America extends far south and deflects 
a part of the eastward moving cold subantarctic 
circumpolar current northward. The resulting 
Peru (or Humboldt) Current is further affected 
by upwelling, so that it has a chilling effect nearly 
to the equator. In the western part of the South 
Pacific, there is no current system as well defined 
as the Kuroshio system. This may be related 
to the great distance separating South America 
and^Australia. According to Sverdrup, Johnson, 

and Fleming (6, p. 706), "in the western South 
Pacific annual variations are so great that in 
many regions the direction of flow becomes 
reversed, as is the case off the east coast of 

Figure 3 shows the current patterns in the 
Atlantic. In the North Atlantic the Gulf Stream 
system, a counterpart of the Kuroshio system in 
the Pacific, is well known for its role in warming 
the shores of both North America (the south- 
eastern part of the United States) and Europe 
(the British Isles and Scandinavia). Among the 
major terminal branches of this system, the 
Irminger Current flows westward to the south of 
Iceland while the Norwegian Current can be 
traced through two further branches into the 
Polar Sea. To the east of Labrador and New- 
foundland, the western part of a counter-clock- 
wise gyral, the cold Labrador Current, is a counter- 
part of the Oyashio Current in the Pacific. Part 
of the North Atlantic Current returns southward 
off the Iberian and North African coasts and 

60 80 100 120 140 160 180 160 140 120 100 80 60 40 20 



<F ~ sb6~"~fooo 

80 100 utud 120^ frrt of 140 0****h 160 160 160 tanjtt*. 140 **< of 12Q o^>w<h 100 

60 <0 

NvtTnoM Scmu or OCM MA 


Fig. 2. Current system and centers of distribution of endemic species of Codium in Pacific Ocean.! Each circle represents 
one species. 





Fig. 3. Current system and centers of distribution of endemic species of Codiwn in Atlantic Ocean. Each circle repre- 
sents one species. 

contributes in part to the North Equatorial 
Current and in part to the Guinea Current, 
which flows along the coast of Africa as far as 
the equator. In the southern hemisphere, the 
shape and position of South America profoundly 
affects the current pattern: the northeastern 
shoulder of Brazil deflects part of the South 
Equatorial Current northward across the equator 
where it flows northwest along the northern 
coast of South America and ultimately contri- 
butes to the Florida Current (the first part of the 
Gulf Stream system). In the Atlantic, unlike the 
Pacific, the surface circulation of the two hemis- 
pheres is thus interconnected. The southern tip 
of South America deflects the cold subantarctic 
circumpolar current not only along the west 
coast, but also along the east coast. The resulting 
Falkland Current meets the southward moving 
branch of the South Equatorial Current (the 


warm Brazil Current) in the latitude of Uruguay, 
where a sharp temperature gradient is thus estab- 
lished. Along the west coast of South Africa, 
the Benguela Current flows northward; and 
being affected by upwelling, it depresses inshore 
temperatures until it gradually leaves the coast 
and continues westward as the South Equatorial 

Much less is known about the current patterns 
of the Indian Ocean (Fig. 4) than of the Pacific 
or Atlantic. A major counter-clockwise gyral 
in the southern part of the Indian Ocean prevails, 
subject to considerable annual variation. The 
South Equatorial Current flows southward around 
Madagascar and along the cast coast of South 
Africa as the warm Agulhas Current. There is a 
well-defined eastward moving current between 
South Africa and Australia, representing the 
returning high-latitude component of the gyral. 


No 00 22 

Fig. 4.--Current system and centers of distribution of endemic species of Codium in Indian Ocean. Each circle represents 
one species. 




On the basis of considerations that temperature 
controls the establishment of populations of 
particular genotypes and that currents provide a 
means of dispersal, we can make predictions con- 
cerning the expected distribution of marine algae. 
In seeking explanations of existing floristic pat- 
terns, however, we must also consider the historical 
factor, despite the difficulty of assessing its role. 
A more recent origin is suggested by taxa whose 
distribution can be explained without recourse to 
the historical factor, while such problems as 
bipolar distribution imply greater antiquity. 

In the Pacific, on the basis of the Kuroshio 
system, we can predict similarities between the 
flora of Japan and that of California. In the At- 
lantic, on the basis of the Gulf Stream system, 
we can predict similarities between the flora of 
the northeastern United States and Atlantic 
Europe. Considering the interconnected surface 
circulation between the northern and southern 
hemispheres of the Atlantic, we might also predict 
similarities between the flora of Brazil and that 
of the Caribbean region. On the basis of the 
Antarctic Circumpolar Current we can predict 
the existence of a circumpolar antarctic or sub- 
antarctic marine algal province. As we shall see, 
each of the four predictions is borne out by data 
from marine algae in general. 

The temperate flora of Japan, although it 
comprises endemic species for the greater part 
and includes a sizable arctic element, is clearly 
related to the California flora. This relationship 
is expressed in similarity of general appearance 
and in the sharing of many genera and even some 
species (including Heterochordaria abietina, End- 
arachne binghamiae, Costaria costata, Cymathaere 
triplicata, Pikea calif arnica, Baylesia plumosa, 
Rhodymenia pertusa, Coeloseira pacifica, Bing- 
hamia californica and Pterosiphonia dipinnatd). 

The similarities between the flora of the north- 
eastern United States and that of Atlantic Europe 
have long been recognized. Among common 
species shared by the two floras, the following 
may be cited: Ulothrix flacca, Prasiola stipitata, 
Cladophora albida, C. flexilis and C. gracilis, 
Bryopsis plumosa, Ectocarpus confer voides, E. 
fasciculatus and E. tomentosus, Giffordia granulosa 
Cladostephus verticillatus, Sphacelaria drrhosa, 
Elachistafucicola, Chordariaflagelliformis, Eudes- 
me virescens, Sphaerotrichia divaricata, Stilo- 
phora rhizodes, Arthrocladia villosa, Desmarestia 
aculeata and D. viridis, Stictyosiphon tortilis, 
Asperococcus echinatus, Desmotrichum undulatum, 

Punctaria plantaginea, Dictyosiphon foeniculaceus, 
Alarm esculent a, Chorda filum and C. tomentosa, 
Laminaria digitata and L. saccharina, Ascophyl- 
lum nodosum, Fucus serratus, F. spiralis and F. 
vesiculosus, Nemalion multifidum, Gelidium crinale, 
Dumontia incrassata, Polyides caprinus, Corallina 
officinalis, Gloiosiphonia capillaris, Cystoclonium 
purpureum, Furcellaria fastigiata, Gymnogongrus 
griffithsiae and G. norvegicus, Phyllophora brodiaei 
and P. membrani folia, Chondrus crispus, Gigartina 
stellata, Rhodymenia palmata, Champia parvula, 
Antithamnion cruciatum, Callithamnion byssoides, 
C. corymbosum and C. roseum, Ceramium diapha- 
num, C. rub rum and C. strictum, Plumaria elegans, 
Ptilota plumosa, Seirospora griffithsiana, Sper- 
mothamnion turneri, Spyridia filamentosa, Mem- 
branoptera alata, Pantoneura angustissima, Phy- 
codrys rubens, Dasya pedicellata, Chondria dasy- 
phylla, Odonthalia dentata, Polysiphonia elongata, 
P. lanosa, P. nigrescens and P. urceolata, Rhodo- 
mela confervoides and R. lycopodioides. 

In searching for similarities between the flora 
of Brazil and that of the Caribbean region, we 
encounter a distributional pattern that is remark- 
able in that it encompasses portions of both 
northern and southern hemispheres. This vast, 
rather homogeneous algal province extends from 
the Caribbean (with some northward extensions 
to Florida, the Carolinas, and Bermuda) to 
southern Brazil and shows certain affinities with 
the flora of the Cape Verde Islands and the 
adjacent African coast and weaker affinities with 
the flora of the Canary Islands and of the western 
Mediterranean. Taylor (8) in a study of 317 
species common in the Caribbean region, found 
that 28.4 % were restricted to the Caribbean, 1 1 .0 % 
extended northward at least to Beaufort, North 
Carolina, 33.1% extended southward along the 
coast of Brazil, and 27.5% extended both north- 
ward and southward. Taylor concludes that 
"the Caribbean flora deserves this name only 
because the Caribbean Sea is the area of its great- 
est known luxuriance and diversity .... [It] is 
actually a west or American Atlantic tropical 
flora which, in spite of the Brazil current and the 
North Equatorial current, extends down the 
Brazilian coast to Rio de Janeiro, with few 
replacements." While the direction of the North 
Equatorial Current would indeed seem to prevent 
or at least to impede the southward spread of a 
Caribbean flora, if the possibility of an eastern 
Atlantic origin be granted, the existence of this 
vast algal province would not be in disagreement 
with hydrographic facts. 

The circumpolar marine algal province en- 
compassing the Antarctic Continent and the 

Subantarctic region (including the southern tip of 
South America and such islands as the South 
Orkneys, South Georgia, South Sandwich, Ker- 
guelen, Heard, Macquarie, Auckland, Campbell, 
and Stewart) has been summarized by Skottsberg 
(5). Characteristic species include Cladophora 
pacifica, Adenocystis utricularis, Caepidium antarc- 
ticum, Desmarestia rossii and D. willii, Durvillea 
antarctica, Geminocarpus geminatus, Halopteris 
funicularis, Macrocystis pynfera, Scytothamnus 
australis and S.fasciculatus, Utriculidium durvillei, 
Balha callitricha, Callophyllis tenera, Chaetangium 
fastigiatum, Delisea pukhra, Lithothamnium an- 
tarcticum, Lophurella hookeriana, Phycodrys quer- 
cifolia, Plocamium hookeri and P. secundatum, and 
Polysiphonia microcarpa. 


To what extent does the distribution of Codium 
agree with previously recognized floristic patterns ? 


First, it should be pointed out that most species 
of Codium are restricted to continuous shores or 
closely spaced islands; only a few species occur 
on two or more widely separated land masses. 
Figs. 2, 3, and 4, indicate the centers of distri- 
bution of endemic species in each of the three 
oceans. The Pacific with 30 endemic species 
leads the Indian with 19 and the Atlantic with 8. 
These numbers undoubtedly will be adjusted 
upward with further study of certain perplexing 
material, as yet unidentified. 

Let us look more closely at the Pacific Ocean. 
Fig. 5 shows the distribution of four closely 
related adherent species. The occurrence of 
C. hubbsii Dawson both in Japan and in California, 
although separate subspecies probably are in- 
volved, should be noted. Codium convolutum 
(Dellow) Silva 2 comb. nov. is an example of a 
local endemic. On the other hand, C. lucasii 
Setchell occurs in South Africa as well as in Austra- 
lia. Codium arabicum Kuetzing is interpreted here 
as a complex series of microspecies or subspecies 

60 80 " 100~ 120 140 160 

180 160 140 120 100 



80 100 lonptudt 12Q fU o( 140 160 180 160 Longitude 14Q Wst of l^Q Gwn^h 1UU HO 


Fig. 5. Distribution of four closely related adherent species of Codium. 

2 Codium adhaerens C.Ag. var. convulotum Dellow, Trans. R. Soc. N.Z. 80: 122. 1952. 



extensively distributed throughout the Indo-Paci- 
fic region, reaching its northeastern limit in the 
Hawaiian Islands. Several interesting patterns 
are shown in Fig. 6. Codium spongiosum occurs 
in South Africa and Mauritius in addition to the 
localities indicated in Fig. 6 and is thus Indo- 
Pacific in distribution. The distribution of each 
of the three members of the highly specialized 
section Digitaliformia is shown: Codium john- 
stonei Silva occurs in California and adjacent 
Mexico, but lacks a Japanese counterpart; C. 
pomoides J. Ag. is restricted to Australia; C. 
dimorphum Svedelius occurs in New Zealand and 
southern Chile. The distribution of each of five 
species of the section Mamilhsa is also shown: 
C. minus (Schmidt) Silva 3 comb. nov. occurs in 
Japan; its nearest relative, C. mamillosum Har- 
vey, occurs in Australia and Hawaii; C. rittcri 
S. and G. occurs in Alaska; the remaining two 
species are distinct from the other members of 
the section and from each other; yet are very 

closely related and are extremely local in distri- 
bution, C. globosum Lucas occurring in Queen- 
sland and C. cranwelliae Setchell in New Zealand. 
Fig. 7 shows the distribution of several dicho- 
tomous species. Codium contractum Kjellman in 
Japan and its very close relative C. macdougalii 
Dawson in the Gulf of California, Mexico, are 
excellent examples of vicarious species. The 
distribution of the three members of the section 
Lata is interesting: Codium latum Suringar, 
described from Japan, is represented by a sub- 
species, C. latum ssp. palmer 7 (Dawson) Silva 4 
comb, nov., on Guadalupe Island, Mexico; C. 
laminar ioides Harvey is endemic to Western 
Australia; while C. platylobium J.E. Areschoug 
(not shown in Fig. 7) is endemic to South Africa. 
Fig. 8 shows the distribution of various repent 
Codium, representing several species or complexes 
of microspecies and subspecies. This group of 
taxa together with the populations of C. arabicum 
indicated in Fig. 5 and C. intertextum Collins and 

60 80 100 120 140 IfaO 180 160 140 120 100 80 60 40 20 


120 . East of 140 Grttnwtch 160 

160 Longitude 140 Wtst of 120 100 

__ Fig. 6. Distribution of Codium spongiosum, of Section Digitaliformia, and of Section Mamilhsa. 

3 Codium mamillosum Harvey var. minus O.C. Schmidt, Bibl. Bot., 23(91): 37, 1923. 

4 Codium palmeri Dawson, Bull. So. Calif. Acad. Sc. % 44: 23, 1945. 



Hervey indicated in Fig. 9 form an integral part of 
the biocoenosis of coral reefs. The anatomical 
variability of these plants from reef to reef is 

Despite the high degree of endemism exhibited 
by Codlum, a study of the distribution of closely 
related species is thus seen to support the recog- 
nition of both the Japanese-Californian floristic 
pattern and the circumpolar subantarctic pattern. 
Moreover, data are provided which support the 
recognition of an Indo-Pacific province, the 
existence of which has already been indicated by 
studies of other algae. This floristic pattern, 
while not strongly suggested by current pat- 
terns, is not unreasonable if migration through 
the Indonesian region is considered possible. 
Finally, an affinity between the floras of eastern 
South Africa and southwestern Australia is 

Now let us turn to the Atlantic Ocean. Fig. 
9 shows the distribution of two closely related 
adherent species, C. adhaerens C. Ag. and C. 
intertextum Coll. and Herv. The populations of 

C. intertextum in the Canary Islands probably 
should be accorded subspecific recognition. Fig. 
10 shows the distribution of another pair of close- 
ly related adherent species, C. effusum (Raf.) 
Delle Chiaje and an undescribed species from 
Tierra d*el Fuego and the Falkland Islands (C. 
antarcticum Silva). This remarkable distribution 
suggests the divergence of a widespread ancestral 
stock into a warm-water and a cold-water species. 
The absence of a closely related species in the 
Caribbean region is notable. Fig. 11 shows the 
distribution of a pair of very closely related 
globose species, C. bursa (L.) C. Ag. and C. 
elisabethae Schmidt, known only from the 
Azores. Again it is interesting to speculate on 
the absence of C. bursa or a closely related species 
in the Caribbean flora. Fig. 12 shows the distri- 
bution of three fairly closely related dichotomous 
species, C. vermilara (Olivi) Delle Chiaje, C 
isthmocladum Vickers, and an undescribed species 
from the Guinea Coast (already in literature as 
C. guineense Silva). The last two species are seen 
to occupy mutually exclusive parts of the tropical 

60 80 100 120'" 140 ~ 160 180 160 140 120 100 80 " "60 40 



80 100 Longnud* 1?Q En o< 140 &w>w.<h 160 180 160 lon t .tuO 140 Weu_* 120 G'tn.>. C h 100 

60 40 

" No DO 21 

Fig. 7. Distribution of two pairs of closely related dichotomous species of Codium. 



Atlantic province. The bipolar distribution of 
C. vermilara is unique and suggests a more 
extensive distribution in former times. Its absence 
in North America is notable. Fig. 13 shows the 
distribution of three distantly related dichoto- 
mous species. Both C. taylori Silva and C. 
decorticatum (Woodw.) Howe occur throughout 
the tropical Atlantic province, although the former 
is absent in the western Mediterranean. Lastly, 
C. tomentosum Stackh., a name at one time applied 
to almost any dichotomous Codium, is seen to 
have a highly restricted distribution. 

It thus can be seen that the temperate North 
Atlantic province is not supported by data from 
Codium because of the absence of this genus on 
the northeastern coast of North America. On 
the other hand, full support is given to the recog- 
nition of the tropical Atlantic province. The 
bipolar distribution of C. vermilara is notable. 


In comparing algal floristic patterns in the 
Pacific with those in the Atlantic and Indian 

oceans, we find that the Japanese-Californian 
alliance is the counterpart of the North Atlantic 
province, but the degree of similarity between 
the Japanese and Californian floras does not 
warrant their recognition as a province. The 
Japanese-Californian alliance is supported by 
data from Codium, the North Atlantic province, 
while lacking such support, is nonetheless well 
documented by distributional data from other 
algae. The tropical Atlantic province has as its 
Pacific counterpart the Indo-Pacific region, which 
not only spans the equator but extends over two 
oceans. Unlike its Atlantic counterpart, it does not 
have affinities with eastern shores; its boundaries 
seem to coincide with the limits of coral reefs, and 
its flora may be an integral part of the coral reef 
biocoenosis. While the components of the flora 
of this vast region are not known sufficiently to 
warrant its designation as a province, it seems 
likely that even after various subregions with 
clusters of endemic species have been removed, 
there will remain a large area with a fairly homo- 
geneous flora. Recognition of both the tropical 

80 100 1ZQ 140 160 

60 140 120 100 60 60 40 20 

No 00 21 

Fig. 8. Distribution of repent Codium. 






No. DD 20 

Fig. 9. Distribution of two closely related adherent species of Codium. 





No. DO 20 



Fig. 10. Distribution of two closely related adherent species of Codium. 




No. DO 20 

A J NrsritoM A Co CHICAGO 

Fig. 1 1 . Distribution of two very closely related globose species of Codiutn. 





Fig. 12. Distribution of three fairly closely related dichotomous species of Codium. 




N O QQ 20 

Fig. 13. Distribution of three distantly related dichotomous species of Codiitm. 



Atlantic province and the Indo Pacific region is 
strongly supported by distributional data from 
Codium. There are no southern hemisphere 
counterparts of the North Atlantic province 
and the Japanese-Californian alliance. The 
circumpolar subantarctic province is well docu- 
mented, including support from Codium. Its 
northern counterpart, the arctic province, al- 
though fully documented (7) has not been consi- 
dered in this paper because of the absence of 
Codium in that part of the world and because 
the existence of this province is dependent upon 
climatic and geographic isolation of a mediter- 
ranean-type sea rather than upon general oceanic 


(1) Bouck,G. B.,and Morgan, E., 1957, The Oc- 

currence of Codium in Long Island Water, 
Bull. Torrey Bot. Club, 84: 384-387. 

(2) Setchell, W.A., 1893, On the Classification 

and Geographical Distribution of the 
Laminariaceae, Trans. Conn. Acad. Arts 
Sr., 9: 333-375. 



__. , 19 15, The Law of Temperature 

Connected with the Distribution of the 
Marine Algae, Ann. Missouri Bot. 
Card., 2 :287-305. 

, 1920, The Temperature Int- 
erval in the Geographical Distribution of 
Marine Algae, Science. 52 : 187-190. 

(5) Skottsberg, C, 1941, Communities of Marine 

Algae in Subantarctic and Antarctic 
Waters, K. Svenska Vetensk-Akad. 
Handl., ser. 3, 19(4), 92 pp. 

(6) Sverdrup,H.U., Johnson, M.W., and Fleming, 

R.H., 1942, The Oceans: Their Physics, 
Chemistry, and General Biology. New 
York, Prentice-Hall, 1087 pp. 

(7) Taylor, W.R., 1954, Cryptogamic Flora of the 

Arctic, II. Algae: Non-planktonic, Bot. 
/fry., 20: 363-399. 

(8) - - ,1 955, Marine Algal Flora of the 

Caribbean Sea and Its Extension into 
Neighboring Seas, in Essays in the Natu- 
ral Sciences in Honor of Captain Allan 
Hancock. Los Angeles, Univ. So.. Calif. 
Press, pp. 259-270. 


M.S. DOTY: Does Codium arabicum occur in southern 

p.c. SILVA: Quite possibly. 

G.F. PAPENFUSS: Very few algae are common to North- 
west Africa and the Caribbean. West Indian species are 
more likely to be found on the east coast of Africa. 

p.c. SILVA: There arc marked temperature differences 
between the coast of Northwest Africa and the Caribbean. 

F.R. FOSBERG: Does Dr. Silva think of these distribu- 
tional patterns of algae as related to currents? Do they 
indicate historical patterns of plant geography, or are they 
related to physiological requirements of the plants? 

p.c. SILVA: The historical factor is very important but 
difficult to assess. A large part of the genus Codium 
seems to be actively speciating although many species arc 
widely separated morphologically and would appear to be 
relicts. The distribution of recent species at least can be 
related to present-day current patterns. 






Fish Culture Research Station, Batu Berendam, Malacca, Federation of Malaya. 

One of the difficulties encountered when read- 
ing past papers on the algae of this part of the 
world is that the areas concerned are not clearly 
defined. Political boundaries rarely coincide with 
biological ones, and the term Malaya has been 
used to cover anything from the whole of the 
Indonesian islands to the Federation of Malaya 
itself. I propose to use it to cover the Federation 
and Singapore, not through any wish to ignore 
the surrounding areas, but because the very 
location of the research station makes it necessary 
that I should confine my main attentions within 
those boundaries. 

Looking through the literature on fresh-water 
algae, one finds that while there are many papers 
dealing with Malay territories generally, very few 
of them deal with the Federation of Malaya and 
Singapore. We have papers by West and West, 
Bernard, Lemmermann, Biswas, and Ruth 
Patrick, but very little else. This is partly because 
in the past botanists in this part of the world 
have been more concerned with the flowering 
plants and ferns, and it is only recently that any 
serious phycological studies have been undertaken. 
With the active interest taken in fresh-water 
biology by departments of the University of 
Malaya, and with the opening of the Fish Culture 
Research Station in Malacca, phycological knowl- 
edge in this area has been considerably advanced, 
although still very inadequate. It is noteworthy 
that the present Acting-Director of the Botanic 
Gardens, Singapore is deeply interested in phy- 
cology, admittedly marine, but still very encourag- 
ing to fresh-water phycologists. 

In a survey of the fresh-water algae of Malaya, 
one might expect certain things, simply by virtue 
of its geography. Firstly it would occasion no 
surprise to find those species which are world- 
wide in distribution, ranging from arctic to tropic 
climates. Superimposed on these distributional 
elements would be found those species which are 
strictly tropical in range, but still circumglobal 
in distribution. Of greater interest, however, 
are those species of much more restricted range. 
The Malay peninsula forms an integral part of a 
geographical system which stretches from Burma 
and Thailand in the north right down through 

the Indonesian islands to Australia. This Indo- 
Malaysian-Australian system was evidently 
almost a continuous land mass at one time. One 
would expect algal species common to the whole 
area, with perhaps minor geographical variations. 
In addition, in Malaya one would expect species 
having greater affinity with those from countries 
in the north, such as India, Burma, and Thailand, 
and other species showing greater similarity to 
those from Java and Sumatra. Finally it is likely 
that a few species may be even more restricted in 
range, being confined almost entirely to Malaya. 

Such distributional patterns will inevitably be 
complicated by the nature of the terrian of the 
country concerned, for the various habitats may 
differ widely in character. Much of the lowland 
of Malaya is lateritic, with swamps and padi 
fields. The water is decidedly acid and desmids 
abound, whilst the common flagellates are various 
species of Chrysophyceae. In many places fish 
ponds have been made, with consequent liming 
and heavy manuring. Euglenineae are dominant, 
or with less heavy manuring, Chlorococcales and 
sometimes Microcystis aeruginosa. On the other 
hand, in the fast flowing streams of the high 
mountains, such as the Cameron Highlands, 
diatoms predominate. The limestone peaks of 
the north have not yet been explored phycolog- 
ically, and should prove to have an interesting 
algal flora. In addition, Malaya is divided cen- 
trally by the main mountain range, and it would 
be interesting to see if those algal species on the 
west side differ in any way from those on the east. 

Despite these distributional complications, 
it should be possible to trace the broad lines due 
to geological and evolutionary history, particular- 
ly if individual groups are studied. Taking the 
very characteristic desmid genus Micrasterias, 
we find those species of known world-wide dis- 
tribution A/, pinnatifida, americana (only once 
recorded in Malaya so far), alata, crux-melitensis, 
thoinasicma var. notata and jenneri. Typical 
circumglobal tropical species or varieties are M. 
mahabuleshwarensis and M. foliacea var. ornata. 
In the Indo-Malaysian-Australian group we have 
M. ceylanica and M. moebii, although the Mala- 
yan form of the latter species is intermediate 



between those from Thailand and Java. M. 
thomasiana var. evoluta and M. apiculata var. 
tjitjeroekensis show greater affinity for forms 
from Java, while M. mahabuleshwarensis var. 
bengalica, M. lux and M. anomala are similar to 
those from India and Burma. So far, M. torreyi 
var. crameri and M. torreyi var. doveri appear 
to be confined to Malaya itself. Further inves- 
tigations may show that some of these species 
are more widespread than at first apparent, but 
the list illustrates the fact that Malaya partakes 
of elements from both northwards and south- 
wards, as well as possessing tropical and world- 
wide elements. Other desmid genera show a 
similar picture, often just as clearly. 

The diatoms of Malaya are not fully worked 
out, and we have insufficient representative 
collections. For that reason, the picture is less 
clear, particularly as so many diatoms are world- 
wide in distribution. Examples of these latter to 
be found in Malaya are Cyclotella meneghiniana, 
Melosira granulata, Synedra ulna, Eunotia pecti- 
nalis, Frustulia vulgar is and Pinnularia braunil. 
So far I have found none which I would call 
purely tropical species, but of much more limited 
distribution are Stenopterobia pelagica and Ach- 
nanthes crenulata, found also in Indonesia, and 
Navicula voigtii occurring also in Thailand and 
S. China. Ruth Patrick mentions one or two 
species which she found only in Malaya, and I 
have found some as yet unnamed ones which 
apparently have not been reported elsewhere, 
but on such limited collections it would be unwise 
to suggest that they are confined to Malaya. 

When we turn to the various flagellate groups, 
we find a wealth of forms of world-wide distri- 
bution also found in Malaya, or else species which 

are recorded for Malaya and for places like 
Germany and France, which suggests they are 
really more widespread. Of the purely tropical 
forms, Phacus pyrum is the most typical, but I 
would not like to be certain about this species, 
for with the flagellates it is often easier to say 
which are the species which will not tolerate 
tropical conditions rather than vice versa. Of 
more limited distribution are Chlamydomonas 
lismorensis, Trachelomonas lismorensis, Trachelo- 
monas volzii var. cylindracea, and Peridinium 
gatunense var. zonatum, all found in both Malaya 
and Australia, while Stromhomonas australica is 
reported from Australia and S. China as well 
as Malaya. It is to be expected that these species 
will be reported from Indonesia as well, Trachelo- 
monas similis var. hyalina, a very characteristic 
species was described from Burma and has proved 
quite common in Malaya. No doubt as more 
work on the flagellates is done in the various 
countries, the picture will become clearer. There 
are several species reported from Indonesia and 
Venezuela, and now from Malaya, and it seems 
possible that they may occur in other tropical 
fresh waters. 

It will be noticed that most of the examples 
I have chosen have well-marked external features, 
so that any differences become obvious. No 
doubt if such groups as the Myxophyceae and 
the Chlorococcales were examined closely, it 
would be found that they show similar distribu- 
tional patterns. 

To sum up, the algal species of Malaya exhibit 
an interaction of worldwide distributional range, 
circumglobal tropical distribution, and a much 
more limited distribution within the Indo-Malay- 
sian-Australian area. 


R.L. CROCKER: You have mentioned geological and 
historical evolutionary trends. Have you any theories 
on this? 

G.A. PROWSE: With the limited amount of information 
available, it would be unwise to formulate any theories. 

O.F. PAPENFUSS: What literary sources are used in 
your work and would Skufa's paper on Burmese algae be 
useful to you ? 

O.A. PROWSE: I have consulted the various libraries 
in Britain and have had photostats made. Good papers 

are available on Indonesia, Burma, and elsewhere, but 
very little on Malaya itself. I have a photostats copy of 
Skufa's paper. 

G.F. pAPfcNFUss: Looking at canals near Bangkok, 1 
would suggest that the Volvocales are common. Is that 
so in Malaya? 

O.A. PROWSE: So far they have been comparatively 
scarce, but possibly may be more frequent in the limestone 






Laboratoire de Biologie Vegdtale Marine, Facttlte de Sciences de Paris, Paris, France. 

The Acrochaetium-Rhodochorton complex, as 
Papenfuss has termed it ( 11 ), groups many little, 
mostly epiphytic red algae, distributed in all the 
oceans. These algae are generally included in 
the order Nemalionales, among which they show 
a more simple structure than the typical forms 
such as Nemalion and Liagora. The vegetative 
fronds consist only of branched filaments with 
heterotrichous habit. 

As I pointed out some years ago (3), this com- 
plex may be distinguished from all other Florideae 
(except perhaps from Gelidiales) because it Jacks 
a true carpogonial branch. In all the Acrochae- 
tiaceae where sexual organs are known, the 
carpogonium is borne directly on the side of a 
normal cell of the filament or at the top of a few- 
celled lateral vegetative branch; sometimes, as 
in Crania, the carpogonium may be intercalary by 
differentiation of a vegetative cell from the erect 

The lack of a carpogonial branch and the ex- 
treme simplicity of the vegetative frond seem to 
me sufficient reasons to put these algae in a 
separate order, the Acrochaetiales. Should this 
order be considered as the more primitive of the 
Florideae or as a reduced group ? 1 am unable to 
settle this definitely. 

The generic classification of the Acrochaetiales 
has for a long time puzzled the phycologists. The 
late Mrs. Drew-Baker (I) united all the species 
in only one genus, the genus Rhodochorton. More 
recently Papenfuss (11 ) proposed a new and much 
more natural classification. He chose, as the 
main distinctive character, the structure of the 
chromatophore and its situation in the cell. 
Unfortunately, shortly before the paper of Papen- 
fuss appeared, Kylin (5) had proposed another 
classification of the Acrochaetiaceae based upon 
morphological characters not as good for a sys- 
tematic classification as those used by Papenfuss. 

Kylin proposed, namely, to place into the genus 
Kylinia, created by Rosen vinge (13) for a micro- 
scopic species epiphytic on Sporochnus in Den- 
mark, all the Acrochaetium species with a single 
basal cell. This character does not seem to me 
a good one. It would be best, perhaps, to take 
into consideration the heterotrichy of these 

plants and, if one would use morphological 
characters for the distinction of genera, to separate 
plants truly heterotrichous from those with only 
erect system originating from a single basal cell 
and those with only creeping filaments. But 
I agree with Papenfuss in considering the type 
of chromatophore as a more important criterion 
for separating genera. 

Kylinia rosulata, type of genus, was described 
by Kylin as having a stellate chromatophore with 
a central pyrcnoid, consequently Papenfuss (12) 
put into this genus all the species previously placed 
by him in his genus Chromastrum characterised 
by its stellate chromatophore. In fact, the plant 
studied by Kylin as Kylinia rosulata is quite 
different from the species described by Rosen- 
vinge. Both species exist on the coast of France 
in the Channel. The species of Kylin is an 
Acrochaetium (Acrochaetium ky Unhides nov. 
sp.) with stellate chromatophore and with only 
asexual monospores. The true Kylinia rosulata pos- 
sesses a parietal chromatophore without pyren- 
oid and its sexual organs are quite different from 
those of Acrochaetium. The male cells (sperma- 
tocysta) are borne at the top of a very long 
hyaline cell that looks like a hair; and Kylin 
thought that Rosenvinge made a mistake, inter- 
preting a normal hair with accidentally attached 
spermatium at the top for the male organs of 
Kylinia. In fact Rosenvinge was right, as I have 
ascertained by the study of living material. 

The carpogonium of Kylinia arises generally 
directly from the spore; and it is, like the vege- 
tative cells, appressed on the filament of the host, 
but the trichogyne is bent upwards. The plant is 
monoecious; and after the fecondation, the 
carpogonium produces directly three carpospores 
at its top. Kylinia rosulata is thus the simplest 
and probably the most reduced type of Florideae 
one can imagine. 

It is particularly noteworthy to recall the 
observations of Levring (7) about a new Kylinia 
he described from Australia, the structure and 
reproduction of which closely recall those of the 
European species. 

The carposporophyte of the other Acrochae- 
tiales is not so reduced, and there are some dif- 



ferences in its development according to the 
different genera. In Acrochaethium (A. borneti, 
A. subtilissimum), the carpogonium is divided, 
after the fecondation, into three or four cells; 
each of them produces lateral branches bearing 
terminal solitary carpospores. In Crania (G. 
effhrescens), the development is the same but the 
carpospores are in chains. 

On the contrary, in Audouinella, as studied by 
Drew (2), the carpogonium does not divide after 
the fecondation but produces directly lateral 
filaments with the carpospores terminal. By 
this character, the fresh-water genus Audouinella 
can be distinguished from the marine genus 

As to the characters of the chromatophores, 
we can distinguish different types among Acro- 

(1) Only one chromatophore per cell; with a 
central pyrenoid. The shape and the 
situation of the chromatophore may be 

a) The chromatophore may be axial in 
the cell with the branches of the chro- 
matophore radiating towards the inner 
surface of the cell in all directions (stel- 
late chromatophore of the genus Chro- 
mastrurn of Papenfuss). 

b) The chromatophore may be parietal 
against the cell wall (genus Acrochae- 
tium s. str. of Papenfuss). This parietal 
chromatophore may be rounded or 
irregularly lobed, sometimes stellate, 
but only in one plane. 

In fact, there arc intermediate forms 
between (a) and (b), and it is difficult, 
I think, to use this character for a 
generic distinction. There will be some 
species, the place of which among 
Acrochaetium or Chromastrum of Papen- 
fuss, will be difficult to settle. 

In some species of this group with 
only parietal chromatophore per cell, 
the pyrenoid is lacking as in Kylinia 

(2) More than one (few to many) parietal 
chromatophores in each cell. 

a) Chromatophores stellate with pyrenoid. 
This is the case for Rhodochorton flori- 
dulum but also for plants up to now 
included in the genus Acrochaetium. 
(A. caespitosum^ A.codii, etc.) 

b) Chromatophores without pyrenoid, 
disc-shaped or more or less spirally 

twisted. This is the case for Rhodo- 
chorton rothii (=R. purpureum) and for 
the genera Audouinella and Crania. 
Considering the mode of development of the 
gonimoblast and the characters of the chroma- 
tophores as well as the probably haplobiontic or 
diplobiontic nature of the life-cycle, one can 
divide the Acrochaetiales into two families and 
eight genera according to the following key: 


Haplobiontic. Only one chromatophore with 
a pyrenoid in each cell. In some species the 
pyrenoid is lacking; in others, there are more than 
one chromatophore per cell, but each is provided 
with a pyrenoid. 

A. Only one chromatophore (central or parietal) 
per cell. 

a) Spermatocysta borne on vegetative un- 
differentiated cells, gonimoblast with spo- 
rogenous filaments bearing terminal car- 

(1) Carpogonium transversally divided 
after fertilization. . . . Acrochaetium 
Naeg. (incl. Colaconema Batt.) 

(2) Carpogonium longitudinally divided 
after fertilization .... Liagoraphila 

b) Spermatocysta borne at the top of dif- 
ferentiated elongated and hyaline cells. 

(1) Carpogonium transversally divided 
after fertilization and producing spor- 
ogenous filaments with terminal car- 
pospores .... Balbiania Sirodot. 

(2) Carpogonium undivided after ferti- 
lization directly producing few car- 
pospores .... Kylinia Rosenvinge. 

B. More than one parietal chromatophore per 
cell, each provided with a pyrenoid .... 
Rhodothamniella nov. gen. 


Probably diplobiontic (or without sexual re- 
production). More than one chromatophore, 
always without pyrenoid, in each cell. 

A. Sexual organs present. Chromatophores 
ribbon-shaped and more or less spirally 
a) Marine Algae. Carpogonium tranversely 

divided after fertilization. Carpospores 

in chains .... Crania Kylin. 



b) Fresh-water algae. Carpogonium un- 
divided after the fertilization. Terminal 
carpospores .... Audoulnella Bory. 
B. Sexual organs unknown. Chromatophores 

disc-shaped .... Rhodochorton Naeg. char. 


From a morphological point of view, the two 
families I propose to distinguish among Acrochae- 
tiales, are very alike as to their vegetative 
structure. But they may not be so closely allied 
as they seem. Cytologically, Acrochaetiaceae 
have a less evolved (more primitive) type of 
chromatophore than the Audouinellaceae. Some 
species of this family (Audouinella) recall the 
Chantransia-stagQ of Batrachospermutn or Lema- 
nea and may be reduced forms. 

Most of our knowledge about Acrochaetiales 
rests upon researches performed on the coasts 
of the Atlantic Ocean. In the Pacific Ocean, 
important work was done upon the species from 
North America by K.M. Drew (1) and upon the 
species from Japan by Nakamura (9,10). More 
recently, some very interesting new species were 
described from Australia and New Zealand by 
Levring (7,8), but Acrochaetiales of large areas 
still remain unknown. 

It is hoped that in the future, accurate studies 
on the morphology as well as the cytology, re- 
production, and life-cycle of Pacific species will 
afford new light upon this most interesting group 
of red algae. 


(1) Drew, K.M., 1928, A Revision of the 

Genera Chantransia, Rhodochorton and 
Acrochaetium, etc. Univ. Calif. Publ. Bot., 
14: 139-124. 

(2) _ , 1935, The Life-History of 

Rhodochorton violaceum (Klitz.), comb. 

nov. (Chantransia violacea Ktitz.), Ann. 
of Dot., 49:439-450. 

(3) Feldmann,!., 1953, devolution desOrganes 

Femelles Chez les Florid^es, Proceed. 
Int. Seaweed Symposium 1952, Edin- 
burgh, 11-12. 

(4) Hamel, G., 1927,Recherches sur les Genres 

Acrochaetium Naeg. et Rhodochorton 
Naeg., Saint-L6. 

(5) Kylin, H., 1944, Die Rhodophyceen der 

Schwedischen Westkiiste, Lunds Univ. 
Arsskr. N.F., Avd 2, Bd. 40, Nr 2. 

(6) ^ , 1956, Die Gattungen der Rho- 

dophyceen, Lund. 

(7) Levring, T., 1953, The Marine Algae of Aus- 

tralia, I. Rhodophyta: Goniotrichales, 
Bangiales and Nemalionales, Arkiv for 
Bot.. Sen 2, Bd 2, Nr 6. 

(8) - 

,1955, Contributions to the Ma- 

rine Algae of New Zealand, I, Arkiv for 
Bot., Ser. 2, Bd 3, Nr 11. 
(9) Nakamura, Y., 1941, The Species of Rhodo- 
chorton from Japan, I, Sc. papers Inst. 
Algol. Research, 2:273-291. 

(10) - ._ _, 1944, The Species Rhodo- 

chorton from Japan, II, ibidem, 3 :99-l\9. 

(11) PapenfussG.F., 1945, Review of Acrochae- 

tium- Rhodochorton Complex of the Red 
Algae, Univ. of Calif. Publ. Bot., 18:229- 

(12) ._ _... _. ,1947, Further Contributions 

toward an Understanding of the Aero- 
chaetium- Rhodochorton Complex of the 
Red Algae, Univ. of Calif. Publ. Bot., 

(13) Rosenvinge, L.K., 1909, The Marine Algae 

of Denmark, I: Rhodophyceae, D. Kgl. 
Danske Vidensk. Sehk. Skrifter. Nat. 
ogMath. Afd., 7, Raekkel(\)\\-\5\. 





Faculty of Fisheries, Hokkaido University, Japan. 

The northwestern part of the Pacific Ocean, 
especially around the northernmost island of the 
Japan Archipelago, or Hokkaido, and the Kurile 
Islands, has been known to be remarkably rich 
in number of genera and species of Laminariales 
since Dr. K. Miyabe, teacher of Tokida, pub- 
lished in 1902 a comprehensive enumeration of 
the kelps in Hokkaido (3). An English edition 
of this paper has recently been prepared by To- 
kida and published by Hokkaido University as 
one volume of the Journal of Faculty of Agricul- 
ture, Hokkaido University (4). In this paper, 
Dr. Miyabe enumerated eight genera, of which 
the genus Laminaria was the largest containing 
as many as twelve species. However, these 
species are not always considered clearly distin- 
guishable from one another by some researchers 
such as Dr. K. Okamura and Dr. S. Ueda, who 
have sometimes expressed their views on the 
basis of their field observations. Nevertheless, 
Miyabe's species have an important practical 
meaning in Hokkaido where most Laminaria 
species are very highly estimated for their econo- 
mic value. Their market value and the scope of 
their use differ remarkably from one another, 
each species representing usually one specific 
locality or habitat. Their local distinctions are 
well reflected in Miyabe's species. 

In recent years, an attempt to transplant a 
Laminaria of superior quality from its home 
locality to other places has been practiced ex- 
perimentally in Hokkaido. In connection with 
this, there arose a question whether such a tran- 
splanted Laminaria was really quite distinct or 
not from other related species of inferior quality. 
If they were nothing but local or ecological forms 
of one and the same species, the transplanted 
Laminaria would not be expected to retain its 
superior quality long enough. On the contrary, 
if it was really a distinct species peculiar to one 
district, its characters would be expected to per- 
sist to some extent. This question may possibly 
be solved not only by the transplanting experiment 
itself, but also by a crossing experiment between 
certain two species. 

Since 1952, cultural experiments of five species 
of Laminaria, viz., L.japonica Aresch., L. ochoten- 

sis Miyabe, L. rellgiosa Miyabe, L. diabolica 
Miyabe, and L. angustata Kjellm., and one species 
of Alaria, viz., A. crassifolia Kjellm., have been 
carried out as follows by us with the object of 
knowing the results of crossing in either couple 
of these species, following the method and 
principle of the study reported by Schreiber about 
twenty years ago, in 1930 (5). 

Zoospores were liberated in autumn or early 
winter. A dilute zoospore suspension was 
pipetted into small Petri dishes containing fil- 
tered sea water. These dishes were placed on the 
table in the room of our laboratory which was 
heated in the daytime by steam from the middle 
of November through the middle of April. The 
gametophytes developed from those zoospores 
remained sterile in such a condition and grew 
within one to two months to minute filamentous 
thalli visible to the naked eye. The growth of 
gametophytes was more rapid in those floating 
on the water surface than in those attached to the 
bottom of the dish. 

Such a filamentous thallus was then picked up 
with a fine platinum needle or a pointed pincette, 
and, after determining under the microscope its 
sex by the size of cells, it was placed in a small 
dish or tube containing Schreiber's culture 

A preliminary cultural experiment of Laminaria 
religiosa under various conditions of temperature 
was carried out from early December, 1952 to 
early February, 1953. The cultural tubes were 
placed in the flasks filled with water and the 
water of each flask was kept respectively at 
8C, 12C, 16C, and 20C constantly all day 
long, and at 20C in the daytime only. Other 
tubes were placed on the table of the room heated 
in the daytime as mentioned above. The growth 
of gametophytes was best of all in the tubes 
kept at 20C only in the daytime and in those at 
room temperature, whereas it was worst in those 
at constant 20C. The isolated gametophytes 
cultured in the tubes at room temperature have 
survived already more than three and one-half 
years and have proved invariably sterile. They 
usually have a minute spherical thallus consisting 

t Presented in abstract by G.F. Parpenfuss. 



of monosiphonous branched filaments. A piece of 
filament taken from these thalli can be used at 
any time as material for a culture experiment, 
and it can grow to a new spherical thallus. On 
the other hand, the gametophytes became fertile 
only when cultured at 8C and 12C, bearing 
sexual organs about 25 days and 30 days after 
the start of culture, respectively, (cf. Fig. 1). 

The eggs thus formed on isolated female 
gametophytes of all the species studied by us 
could germinate and develop parthenogenetically. 
The development of those parthenosporophytes 
was in general more or less irregular and resulted 
in the production of variously shaped malformed 
thalli as illustrated in Figs. 2 and 3. Malformed 
sporophytes of similar structure were once re- 
ported by Kanda (2) to occur in a mixed culture 
of Lamlnaria ochotensis Miyabe (2, Fig. 4, 9-10), 
L. cichorioides Miyabe (Fig. 7, 2-1 1) and L. sacha- 

linensis Miyabe (Fig. 10, 4). In view of the very 
common occurrence of parthenogenesis in our 
culture experiments, resulting in the production 
of malformed thalli and the occasional produc- 
tion of malformed sporophytes in Kanda's mixed 
culture, the presence in nature of parthenoge- 
netically developed malformed haploid embryos 
of kelps are supposed to be not improbable. 
However, malformed adult thalli of kelps, such as 
lately reported from Hokkaido and vicinity (cf. 
/, 6, 7, 8) are not only of very rare occurrence 
but also are considered not to have been derived 
from such embryos as mentioned above. Such 
haploid embryos of kelps may possibly be of a 
limited life span. 

Crossing experiments were carried out by 
putting in a tube a female or male gametophyte 
of one species together with a male or female of 
another which had been cultured in isolation. 
From twenty to twenty-two tubes were prepared 

Fig. 1 . Growth rate of gametophytes of Laminaria religiosa as indicated by the number of body cells and time of the first 
appearance of reproductive organs (denoted by x) in relation to the water temperature. 



Fig. 2. Showing various forms of malformed parthenosporophytes of Laminar ia religiosa in various stages of development. 
(Magnification: x 200, x 170, x 95, and x 20.) 




Fig. 3. Showing various forms of malformed parthenosporophytes of the following four species of Laminaria. 1-5, 
Lamina riajaponica;6-S,L. ochotensis;9, L. angustata\ 10-11, L.diabolica (Magnification: x 200, x 170, x 95, and x 20). 






Fig. 5. Microphotographs of embryonal Laminar/a sporophytes produced in the following six crossing experiments. 
A, Laminaria japonica 9 X L. ochotensis c?, x 50; B, Laminaria religiosa 9 X L.japonica <, x 50; C, Laminaria ochotensis 9 
X L. religiosa cT, x 80; D, Laminaria religiosa 9 X L. ochotensis <J, x 80; E, Laminaria angustata 9 X L. religiosa #, x 50; 
F, Laminaria ochotensis 9 X Alaria crassifolia <j, x 120. 

Fig. 4. Microphotographs of Laminaria gametophytes in various stages of development. A-H, J-N, R, U, Laminaria 
religiosa', A, one month old gametophytes cultured at constant 20C; B, twenty-five days old female gametophytes bearing 
eggs in a culture kept at constant 8C; C, sixty days old gametophytes in a culture kept at 20C only in the daytime; D, 
seventy-seven days old female gametophyte bearing a parthenosporophyte, floating on the surface of cultural solution; E, 
thirty days old gametophytes cultured at constant 16C; F, seventy-seven days old female gametophyte floating on the 
surface; G, seventy-seven days old female gametophyte attached to the bottom of the cultural dish; H, seven months old 
male gametophyte floating on the surface; J-N, five stages of development of one and the same piece of filament which was 
first taken at the stage shown in J from an immature female gametophyte floating on the surface; K, two months old; L, 
two and one half months old; M, three months old; N, four and one half months old; R, an immature female gametophyte 
cultured in isolation, 3.7 months old; U, a mature female gametophyte grown from a thallus similar to that which is illus- 
trated in R after placed in a newly prepared Schreiber's solution three weeks before. I, Laminaria japonica, three months old 
gametophyte floating on the surface. O-P, Laminaria angustata; O, female and male gametophytes in a nine days old 
culture, at this stage of growth, isolation of gametophytes was done; P, a female gametophyte isolated 121 days before. 
Q,T, Laminaria diabolical Q, a female gametophyte bearing parthenosporophytes, isolated 78 days before; T, a female 
gametophyte bearing parthenosporophytes, 68 days after isolation. S, Laminaria ochotensis, a female gametophyte 3.5 
months old, bearing abundant parthenosporophytes. (Magnification: A, E-H, J-M, O-R, U x 80; B x 320; C-D, I, N x 40; 
S-Tx 120). 



for each crossing pair. The result of the experi- 
ments are summarized in Table 1. 

Table 1. 

Result of crossing experiments, showing the 
combination of species crossed and the structure, 
whether normal or abnormal, of sporophytes 

Couple of species crossed 





L. japonica & L. religiosa 
L. japonica & L. ochotensis 
L. religiosa & L. ochotensis 
L. religiosa & L. angustata 
L. ochotensis & L. angustata 
L. japonica & A. crass if olia 
L. religiosa & A. crass if olia 
L. ochotensis & A. crass if olia 





The mark + in the Column "Normal" of Table 
1 denotes that a larger number of the sporophytes 
produced in the corresponding couple of species 
are normal in shape. The parent species of these 
normal sporophytes are supposed to have achieved 
interspecific fertilization. Such a positive result 
of a crossing experiment in Laminaria is believed 
to be new to science, provided that the three 
species concerned are all valid. Or, this result 
may be taken rather as an evidence of specific 
identity of the three species. However, in our 
present state of knowledge, any conclusive remark 
is to be reserved until more decisive data, cytolo- 
gical and ecological, are acquired. 


(I) Hasegawa, Y. et #/., 1956, Notes on the Ab- 
normal Laminaria Fronds Found in Hok- 

kaido and Aomori Pref. (In Japanese), 
Monthly Rept. Hokkaido Fish. Exp. 
St., 13: 69-70. 

(2) Kanda, T., 1946, Culture Studies of Lamina- 

riaceous Plants in Hokkaido, (In Ja- 
panese), Jour. Inst. Res. Sc. Fish., 
Hakodate Coll. Fish., No. 1, 1-44. 

(3) Miyabe, K., 1902, The Laminariaceae of 

Hokkaido, (In Japanese), Rept. Invest. 
Mar. Resour., Hokkaido, 3: 1-60. 

(4) , 1957, The Laminariaceae of 

Hokkaido (1902). An English edition, 
translated by J. Tokida, Jour. Fac. Agr. 
Hokk. Univ., 1, (In the press). 

(5) Schreibcr, E., 1930, Untersuchungen iiber 

Parthenogenesis, Geschlechtsbestim- 

mung und Bastardierungsvermogen bei 
Laminarien, Planta, 12 : 331-353. 

(6) Tokida, J. et al, 1956, On Two Rare Abnormal 

Forms of Laminaria japonica Aresch. 
(In Japanese), Collecting and Breeding, 
18: 118-119. 

(7) , 1956a, On a Malformed La- 

minaria with a Spirally Twisted Lamina, 
(In Japanese), Monthly Rept. Hokk. 
Fish. Exp. St., 13 : 408-411. 

(8) , 1957, On Several Examples 

of Malformed Kelps in Hokkaido and 
Vicinity, (MS read before the 10th 
Hokkaido Regional Conference of the 
Botanical Society of Japan, 29-30 June 

(9) , 1957a,AChimaeraof,4/tfr/a 

and Laminaria Found in Nature, (MS 
sent to the editors of "Nature"). 


G.F. PAPENFUSS apologized for summarizing the paper 
in the author's absence, and regretted that the illustrations 
had not arrived. 

R.F. SCAGEL: How were the cultures started? 

G.F. PAPENFUSS (consulting original script): In petri 
dishes, the floating gametophytes being isolated to flasks 
and maintained at room temperature. Sub-culturing was 
carried out from these gametophytes. 

R.F. SCAGEL: Care must be taken when drawing con- 
clusions from such techniques. All gametophytes look 
alike (apart from the cell differences in male and female), 
and the sporophyte must show secondary morphological 
features before the species can be identified. 

In reply to Dr. Papenfuss' enquiry, Dr. Scagel said that 
his laboratory isolated sporangia to start their cultures. 






Department of Botany, University of California, Berkeley, California, U.S.A. 

There is ample reason to believe that the current 
interest in the algae as research material for many 
kinds of problems in biology and biochemistry 
and as possible sources of food and fuel will 
continue. This interest has had and will continue 
to have a stimulating effect on those aspects of 
phycology that are primarily concerned with the 
acquisition of basic knowledge about the algae 
for its own sake. We may look forward, there- 
fore, to expanded opportunities for fundamental 
work on these plants and to an increase in the 
number of phycologists in the world. 

Because of its large size and the countless 
variety of habitats that it provides, no other ocean 
possesses such a rich and diversiiied flora as the 
Pacific. Our knowledge of the algae of large parts 
of the Pacific is still extremely meager, however. 
Many of the genera and species have been col- 
lected only once and are known only from the 
brief and often inadequate original descriptions. 
The flora of the Pacific thus provides us with 
unlimited possibilities for the pursuit of basic 
problems in all branches of phycology. 

In anticipation of the development of oppor- 
tunities for work on the algae of areas away from 
the present centers of activity in the Pacific, I 
have selected for discussion certain problems 
relating to the taxonomy of green, brown, and 
red algae occurring in some of the more remote 
parts of this ocean. The problems chosen are 
of interest because their solution not only will 
advance our knowledge of a number of little- 
known genera, some of which are also represented 
in the floras of other parts of the world, but the 
information gained will be useful in the inter- 
pretation of natural relationships within the 
groups concerned. 


Information is needed on the life history, the 
morphology of the flagella of the motile cells, 
the kinds of pigments present in the plastids, and 
the nature of the food reserves of Pseudodicho- 
tomosiphon in order to determine whether this 
genus belongs to the chlorophycean family Dicho- 
tomosiphonaceae, which was established by Feld- 
mann (12) for the fresh-water genus Dichotomosi- 

phon, or whether it is a member of the Vaucheria- 
ceae, and hence representative of the class Xan- 
thophyceae of the phylum Chrysophycophyta. 
Pseudodichotomosiphon is a monotypic genus that 
was established by Yamada in 1934 upon material 
from the Ryukyu Islands. Tseng (33) has obtained 
it also on the Island of Hainan. Luther (20, p. 
36) is of the opinion that P. constricta (Yamada) 
Yamada had best be retained in Vaucheria. 

On the basis of material from Taiwan, Hey- 
drich in 1894 erected the genus Rhipidiphyllon. 
He regarded the type material as representative 
of the species which Askenasy in 1888 described 
under the name Anadyomene reticuhita upon 
material from Western Australia. More recently 
B^rgesen (4) and Taylor (31 ) have reported R. 
reticulatum from Easter Island and the Marshall 
Islands, respectively. Two problems relating to 
this genus are: (1) Is the type material actually 
representative of the plant from Western Austra- 
lia? (2) Is Rhipidiphyllon an autonomous genus 
or should it be merged in Microdictyon, with 
which it appears to agree in all essential details? 

Rhipidodesmis A. et E.S. Gepp (13), Boodleopsis 
A. et E.S. Gepp (13) and Pseudochlorodesmis 
B^rgesen (4) are three genera accredited to the 
Codiaceae that perhaps are merely the filament- 
ous, unconsolidated juvenile stages of other 
genera of the Codiaceae, such as Udotea or 
Avrainvillea, which are well represented in the 
Pacific, or Penicillus, which has not yet been 
reported from the Pacific. In this connection, 
I might remark that I obtained in South Africa 
filamentous, unconsolidated mats of Codium 
that are easily mistaken for a filamentous genus 
of the Codiaceae; in fact, they were so mistaken 
by Dr. Egerod and me until Dr. Silva assured us 
that they were only the juvenile stages of some 
species of Codium. 

Rhipidodesmis is based upon a species from 
Ceylon, R. caespitosa (J. Agardh) A. et E.S. 
Gepp, which has also been obtained on the 
Island of Hainan by Tseng (33). Boodleopsis was 
erected for a species from the Malayan Archi- 
pelago, B. siphonacea A. et E.S. Gepp, and 
Pseudochlorodesmis is based upon a plant from 
the Canary Islands, P. furcellata (Zanardini) 

t The preparation of this paper was aided by a Grant-in-aid from the National Science Foundation. 



B^rgesen. This species was recently reported 
from Vietnam and the Marshall Islands by 
Dawson (7,9). 

In the paper that I gave at the Seventh Pacific 
Science Congress in New Zealand (23), I have 
already remarked upon the uncertain status of 
Rudicularia (? Codiaceae), which was described 
by Heydrich (16) from material obtained in the 
Ryukyu Islands, and of Bryobesia (? Derbesia- 
ceae) which Weber-van Bosse (34) founded for 
a plant from Java. 


Troll in 1931 (32) described under the generic 
name Dictyotopsis a peculiar monostromatic 
plant which grows by means of wedge-shaped 
apical cells that cut off segments alternately on 
two sides. The plant forms dense growths on 
the aerial roots of mangroves in river mouths in 
Sumatra and Amboina. Reproductive organs, 
other than propagules, were not observed in any 
of the material. In a general way, Dictyotopsis 
bears some resemblance to the liverwort genus 
Metzgeria, but it differs from this genus in a 
number of important features. Dictyotopsis is 
provisionally placed in the order Dictyotales, 
but the type of apical cell in its thallus is quite 
unlike that characteristic of the thalli of the 
Dictyotales. Pigment analysis and the discovery 
of reproductive organs will probably settle the 
question of the systematic position of this inter- 
esting genus. 


At the generic level, the red algae of the Pacific 
present more problems than any of the other large 
groups of marine algae. The following are some 
of the genera that would especially repay study. 

Gracilariocolax Weber- van Bosse (38), which 
is based on a species that occurs as a parasite on 
Gracilaria in Java, is considered a genus of doubt- 
ful systematic position by Kylin (17). It would be 
instructive to make a comparative study of 
Gracilariocolax, Gracilariophila Setchell et Wilson 
(39), which is a parasite on Gracilaria along the 
Pacific coast of North America, and Holmsella 
Sturch (28), which is a parasite on Gracilaria in 
England, to see whether there is a relationship 
between these three genera. None of them has 
been thoroughly studied, and it is conceivable 
that they will be found to be closely related, if 
not congeneric. 

i De Toni fil. (10) has proposed the substitute name Drouetia for this genus. 

The genus Weberella (Rhodymeniales: Rhody- 
meniaceae) was described by Schmitz (25) upon 
material from Flores Island in the Malayan 
Archipelago. The type and only species, W. 
micans, has also been reported from Timor and 
Java by Weber-van Bosse (38) and from Taiwan 
by Yamada (40). A study of the structure and 
reproduction of this genus is highly desirable. 
To judge from the original description by Schmitz 
and the observations of Yamada, it appears likely 
that Weberella and Herpophyllon Farlow 1 (11), 
which is based upon a type from the Galapagos 
Islands, are congeneric. Kylin (17) has placed 
Herpophyllon in the Squamariaceae (Cryptone- 
mialcs), but the structure of the thallus and es- 
pecially the habit of these two genera (both have 
a dorsiventral organization and produce hap- 
teroid protuberances on the lower surface of the 
thallus whereby the lobes of the thallus become 
secondarily attached to one another or to the 
substratum) suggest that they belong in the 
Rhodymeniales. In connection with these two 
genera, attention should also be drawn to Mali- 
chrysis (J. Agardh) Schmitz (24), which is based 
upon a type from Morocco, and Sciadophycus 
Dawson (6), which was founded for a plant from 
Cerros Island (off the west coast of Baja Califor- 
nia), both of which appear to belong to the same 
complex as Weberella and Herpophyllon. 

On the basis of material from the Malayan 
Archipelago, Weber-van Bosse (34) established 
the genus Exophyllum. This plant has been re- 
ferred, with doubt, to the Rhodymeniaceae. 
One of its characteristic features is the production 
of tetrasporangia in stichidia, a character which, 
as Kylin (17, p. 558) has remarked, is entirely 
foreign to the Rhodymeniaceae or for that matter 
to the order Rhodymeniales. The determination 
of the systematic position of this plant must 
await a study of its reproductive organs. Recent- 
ly the genus was discovered in southern Japan 
by Tanaka (30). 

Zellera and Opephyllwn are two of the little- 
known genera of the Delesseriaceae. Zellera 
was described by Martens in 1866 upon a plant 
from the Island of Tawalli in the Moluccas and 
was later reported from the same region by 
Weber-van Bosse (36). The genus appears to be 
related to the reticulate genera Claudea and 
Vanvoorstia, but its true affinities will remain 
uncertain until well-preserved material of the 
type species, Z. tawallina, has been studied. 
Martens stated that the blades tend to fuse with 
others to form a reticulum; but this is not the 



case, according to Sluiter (27) and Weber-van 
Bosse (36). Knowledge of the structure and 
reproduction of Z. tawallina may also assist in 
the solution of a problem relating to the algae of 
the West Indies. Sluiter (27) has referred a species 
from that region to Zellera, but her figure of its 
habit gives the impression that the plant is mis- 
placed in Zellera. 

Opephyllum was erected by Schmitz (25) for 
a species collected by Martens at Zamboanga 
on the Island of Mindanao. This plant has never 
again been collected. The genus has not been 
illustrated; but to judge from Schmitz's descrip- 
tion, it seems likely that Opephyllum will be found 
to be congeneric with Martensia Hering. 

As is well known, Madame Weber-van Bosse 
described a relatively large number of new genera 
and species from the East Indies. Unfortunately, 
she did not always furnish the detailed informa- 
tion that is necessary for the correct placement 
of her taxa. The following are genera of red 
algae established by Weber-van Bosse, in addition 
to some of those already referred to, whose 
systematic position is uncertain: Dorella (1921), 
Catenellocolax (1928), Microphyllum (1928), 
Corallophila (1923), Mortensenia (1926), Chali- 
costroma (1911), and Perinema (191 1). 

To illustrate the benefits for systematic phy- 
cology to be derived from a study of the old 
taxa, I might review briefly a few of the recent 
contributions that have materially advanced our 
knowledge of certain genera. 

In the paper that 1 gave at the Seventh Pacific 
Science Congress (23), I remarked that the occur- 
rence of the reproductive organs on filaments in 
the New Zealand genus Microzonia is different 
from the condition in all other Dictyotales, to 
which order this genus had been assigned. A 
study by Miss O'Donnell (22) of material of this 
genus given me by Lindauer has shown not only 
that the reproductive organs (unilocular spor- 
angia) actually are produced on filaments but 
that the thallus shows trichothallic growth. It is 
evident, therefore, that Microzonia does not 
belong to the Dictyotales but to the Cutleriales, 
which order previously comprised only the 
genera Cutleria and Zanardinia. 

Levring (18,19, and in Svedelius, 29) studied 
newly-collected material as well as the type of the 
type species of Gloiophloea, G. scinaioides J. 
Agardh (2), and found that our concept of this 
genus is wrong. The previous concept was that 
developed by Setchell (26) in his monograph on 
the Scinaia complex and was based on a plant 
from New Zealand which Setchell believed to be 

representative of G. scinaioides, whose type (not 
seen by Setchell) was obtained in Australia. 
Levring established that the species from New 
Zealand (and also the other species of Gloiophloea 
described by Setchell) is representative of a new 
genus for which he proposed the name Pseudo- 

The genus Corallopsis was erected by Greville 
in 1830 for Sphaerococcus salicornia C. Agardh 
(1), whose type was collected by Chamisso during 
the voyage of the Russian exploration ship 
Rurik, allegedly in Unalaska. Several authors 
have doubted that Unalaska actually was the 
source of Chamisso's material. Dawson (8) 
recently collected a plant in the Philippines that 
agrees in all essential details with C. Agardh's 
(I) illustration of the type of C. salicornia. 
Examination of the log of the Rurik revealed, 
furthermore, that the ship had stopped for six 
weeks at Manila, which circumstance suggests 
with a good deal of certainty that the type of 
C. salicornia was obtained in the Philippines. 
Dawson's study of the material that he collected 
in the Philippines has shown that C. salicornia 
is a species of Gracilaria. Thus Dawson in a 
most satisfactory way cleared up the question 
about the source of the type of C. salicornia and 
at the same time showed that Corallopsis Greville 
(14) is congeneric with Gracilaria Greville (14). 

From the examples that have been discussed 
of problems awaiting investigation and of prob- 
lems solved, it is evident that the flora of the 
Pacific offers challenging opportunities for fund- 
amental work in systematic phycology. 

I have confined my discussion to problems 
relating to the type species of genera occurring 
in more or less remote parts of the Pacific. In 
addition, many taxa from such regions have been 
described as new species of old genera and others 
have been identified with species based on types 
from other parts of the world. Detailed study of 
these species will reveal that a large number of the 
putative new species actually are not new and that 
many of the others have been misidentified. 
Therefore, on the basis of present knowledge of 
the algae of the Pacific, little can be concluded 
with any degree of certainty about the geographic 
distribution of the majority of the species in this 
large ocean. 

I do not wish to convey the impression that in 
my opinion nothing is to be gained from the 
cataloguing of the floras of unexplored areas. 
It is true, however, that in such works further 
confusion will be created through the inevitable 
description of new taxa, many of which will have 



to be reduced when the old taxa have been care- 
fully studied. 

I should thus like to recommend to the phyco- 
logist for whom an opportunity has developed to 
work in remote parts of the Pacific, that he selects 
as territory of operation one or more of the 
regions where previous collecting has been done 
rather than an area that has remained unexplored. 
The unexplored parts can be taken care of much 
more profitably after the forest of confusion has 
been removed from the old trails. 


(1) Agardh,C. A., 1 820Jcones Algarum Ineditae, 

fasc., 1 (4) pp., 10 pis., Lund. 

(2) Agardh, J.G., 1872, Bidragtijl Florideernes 

Systematik, Lunds Univ. Arsskr. Afd., 2, 
8(6), 60 pp. 

(3) Askenasy, E., 1888, Algen mit Unterstiit- 

zung der Herren E. Bornet, A. Grunow, 
P. Hariot, M. Moebius, O. Nordstedt 
bearbeitet, in A. Engler (Ed.), For- 
schungsreise S.M.S. "Gazelle", Theil 4, 
Botanik, pp. 1-58, pis. 1-12, Berlin. 

(4) B^rgesen, F., 1920, Marine Algae from 

Easter Island, in C. Skottsberg, The 

Natural History of Juan Fernandez 

and Easter Island, 2:247-309, 50 figs., 

(5) , 1925, Marine Algae from the 

Canary Islands ... 1: Chlorophyceae. 

K. Danske Vidensk. Selskab, Biol. Medd. 

5(3), 123 pp., 49 figs. 
(6) Dawson, E.Y., 1944, Some New and Unre- 

ported Sublittoral Algae from Cerros 

Island, Mexico, Bull. So. Calif. Acad. 

Sci., 43:102-112, 20 figs. 

1954a, Marine Plants in the 


(8) -_- - 

Vicinity of the Institut Oceanographique 
de Nha Trang, Viet Nam, Pacific Sci., 
8:373-469, 63 figs., 1 map. 

, 1954b, Notes on Tropical 
Pacific Marine Algae, Bull. So. Calif. 
Acad. Sci., 53:1-7, 4 figs. 

, 1956, Some Marine Algae of 

the Southern Marshall Islands, Pacific 
Sci., 10:25-66, 66 figs. 

(10) De Toni, J., 1938, A Note on Phycological 

Nomenclature, Rhodora, 40:27. 

(11) Farlow, W.G., 1902, Thallophytes and 

Musci of the Galapagos Islands, Proc. 
Amer. Acad. Arts Sci., 38:82-99, 102-104. 


(12) Feldmann, J., 1946, Sur I'h6t6roplastie de 

Certaines Siphonales et leur Classifica- 
tion, C.R. Acad. Sci. (Paris), 222:752- 

(13) Gepp, A., and Gepp, Ethel S., 1911, The 

Codiaceae of the Siboga Expedition . . . 
Siboga-Exped. Monogr., 62, 150 pp., 22 
pis., Leiden. 

(14) Greville, R.K., 1830, Algae Britannicae, ... 

88 + 218 pp., 19 pis., Edinburgh. 

(15) Heydrich, F., 1894, Beitrage zur Kenntniss 

der Algenflora von Ost-Asien Besonders 
der Insel Formosa, Molukken-und Liu- 
kiu-Inseln, Hedwigia, 33: 267-306, pis. 
14, 15. 

(16) , 1903, Rudicularia, ein Neues 

Genus der Valoniaceen, Flora, 92:97- 
101, 4 figs. 

(17) Kylin, H., 1956, Die Gattungen der Rhodo- 

phyceen, 15 + 673 pp., 458 figs., Lund. 

(18) Levring, T., 1953, The Marine Algae of Aus- 

tralia, I. Rhodophyta: Goniotrichales, 
Bangiales and Nemalionales, Arkiv Bot. 
ser. 2,2:457-530, 55 figs. 

(19) , 1956, Contributions to the 

Marine Algae of New Zealand, T. Rhodo- 
phyta: Goniotrichales, Bangiales, Nema- 
lionales and Bonnemaisoniales, Arkiv 
Bot. ser. 2, 3:407-432, 15 figs. 

(20) Luther, H., 1953, Vaucheria schleicheri de 

Wild, neu ftir Nordeuropa, Mem. Soc. 
Fauna Flora Fennica, 28:32-40, 5 figs. 

(21) Martens, G., 1 866, Die Tange. Preuss. Exped. 

Ost-Asien, Bot. Theil., (1) + 152pp., 8 pis., 

(22) O'Donnel, Elsa H.J., 1954, The Structure 

and Taxonomic Position of the Brown 
Alga Microzonia, Amer. Jour. Bot., 41: 
380-384, 12 figs. 

(23) Papenfuss, G.F., 1953, Outstanding Prob- 

lems in the Morphology and Taxonomy 
of the Marine Algae of the Aropical and 
Southern Pacific, Proc. Seventh Pacific 
Sci. Congress, 5, Botany, pp. 27-39. 

(24) Schmitz, C.J.F., 1889, Systematische Cber- 

sicht der bisher Bekannten Gattungen 
der Florisdeen, Flora, 72:435-456, pi. 21. 

(25) Schmitz, C.J.F., and Hauptfleisch, P., 

1896-1897, Rhodophyceae, in A. Engler 
and K. Prantl, Die natiirlichen Pflanzen- 
familien, Tell 1, Abt., 2, pp. 298-544, figs. 
192-288, Leipzig. 

(26) Setchell, W.A., 1914, The Scinaia Assem- 

blage, Univ. Calif. Publ. Bot., 6:79-153, 
incl. pis. 10-16. 

(27) Sluitcr, Catharina P., 1908, List of the Algae 

Collected by the Fishery-Inspection Cu- 
racao, Rccueil Trav. Bot. Norland, 4: 
231-241, 1 fig., pi. 8. 

(28) Sturch, H.H., 1926, Choreocolax Polysipho- 

niac Reinsch., Ann. Bot., 40:585-605, 
15 figs. 

(29) Svedelius, N., 1956, Are the Haplobiontic 

Florideae to be Considered Reduced 
Types? Sv. Bot. Tidskr., 50: 1-24, 14 figs. 

(30) Tanaka, T., 1950, Studies on Some Marine 

Algae from Southern Japan, 1. Jour. 
Kagoshima Fish. College, 1:173-180, 
4 figs., 1 pi. 

(31) Taylor, W.R., 1950, Plants of Bikini and 

Other Northern Marshall Islands, 15 + 
227 pp., Frontispiece, pis. 1-79, Ann 

(32) Troll, W., 1931, Botanische Mitteilungen 

aus den Tropcn, IIT: Dictyotopsis pro- 
pagulifera W. Troll, eine nene Brack- 
wasseralge Ostindischer Mangrovege- 
biete, Flora, 125:474-502, 18 figs., 1 

(33) Tseng, C.K., 1936, Studies of the Marine 

Chlorophyceae from Hainan, Amov 
Marine Blot. Bull., 1:129-200, 34 figs*., 

(34) Weber-van Bosse, Anna, 191 1, Notice sur 

quelques Genres Nouveaux d'algues de 


PArchipel Malaisien, Ann. Jar din Bot. 
Buitenzorg, ser. 2, 9:25-33. 

(35) ----- - -, 1921, Liste des 

Algues du Siboga, H: Rhodophyceae, 
partie 1, Protofiorideae, Nemalionales, 
Cryptonemiales, Siboga-Exped. Monogr. 
596, pp. 185-310, figs. 53-109, pis. 6-8, 

(36) - -- , 1923, Liste des 

Algues du Siboga, III: Rhodophyceae, 
partie 2, Ceramiales. Siboga-Exped. 
Monogr. 59c\ pp. 311-392, figs. 110-142, 
pis. 9, 10, Leiden. 

(37) , 1926, Algues de 
['expedition danoise aux iles Kei. Viden- 
skab. Medd. Dansk Naturhist. For. 
Kfbenhavn, 81:57-155, 43 figs. 

(38) - , 1928, Liste des 
Algues du Siboga, IV: Rhodophyceae, 
partie 3, Gigartinales et Rhodymeniales, 
Siboga-Exped. Monogr. 59d, pp. 393- 
533,71 figs., pis. 11-16, Leiden. 

(39) Wilson, Harriet L., 1910, Gracilariophila, 

a New Parasite on Gracllaria confer- 
voides, Univ. Calif. Publ. Bot., 4:75-85, 
incl. pis. 12, 13. 

(40) Yamada, Y., 1932, Notes on Some Japanese 

Algae IV, Jour. Fac. Sci. Hokkaido Univ. 
ser. 5, 2:267-276, 3 figs., pis. 3-9. 
(41) , 1934, The Marine Chlorophy- 
ceae from Ryukyu, Especially from the 
Vicinity of Nawa, Jour. Fac. Sci. Hokkai- 
do Univ. ser. 5, 3:33-88, 55 figs. 


M.S. DOTY: I suggest that the algal subcommittee 
compile a list of problematic species and their localities. 

G.F. PAPENFUSS: It is vital to clear up the confusion in 
old species before erecting new ones. 

j. FELDMANN: Few species are completely well-known. 
More cytological studies should be carried out on genera 
such as Vaucheria. 

G.F. PAPFNFUSS: The problem becomes simpler if there 
is a plentiful supply of material. 

G.A. PROWSE: In the desmids, the frequent occurrence 
of dichotypical cells has made the rc-asscssment of the 
concept of species vitally essential. 

M.S. DOTY: Have the pigments of Dichotomosiphan 
been investigated? 

j. FELDMANN: On the basis of the plastids, there is no 
doubt that it is a green alga. 

G.F. PAPENFUSS: Dr. Feldmann is fully justified in 
erecting the family Dichotomosiphonaceae. 






Department of Botany, University of the Philippines, Quezon City, Philippines. 

One finds the first general attempt to identify 
the algae of the Philippines in Blanco's Flora 
dc Filipinos, of which there were three editions, 
in 1837, 1845, and in 1877 to 1883, the latter 
edition posthumous. Blanco's identifications of 
Philippine species with those of older authors 
were of necessity unreliable as he worked in 
botanical isolation and with very little available 
literature. He proposed a few names for new 
species himself in his first edition, of 1837, which 
arc no more indefinite that some of those of his 
contemporaries in other parts of the world. 
These names were later changed (in one instant 
by himself) or were considered by other botanists 
as synonyms of older ones. In his second edition, 
Blanco himself replaced the name Fucus Gidaman 
Blanco by Fucus edulis, which was considered by 
Georg von Martens to be Sphacrococcus gelatinus 
Agardh. The latter name was based upon 
Euchcuma gclatinae and Fucus gelatinus.i 

Von Martens made a serious attempt to syno- 
nymizc Blanco's algae and proposed the following 

Fucus prolifcr = Halimeda discoidea Decaisne 
Fucus denticulatus - Sargassum (spp.) 
Fuc us Gulaman - Sphacrococcus gelatinus Ag. 
An unnamed species similar to last - Sphaero- 
carpus liclicnoides Ag. 

Viva umbilicalis = Zonaria gymnospora Kiitzing 
Viva compressa ^~ Enteromorpha complanata, 

var. crinita Kilt zing 
Viva intcstinalis - Enteromorpha intcstinalis 


It must be noted that some of Blanco's names 
are apparently older than the synonyms, and if 
sufficiently definite in application to be synony- 
mi/cd, should probably replace the later names. 
This is particularly true as Blanco was rather 
precise in indicating localities, where new collec- 
tions at the places he indicated might enable his 
species to be interpreted. This is a matter that 

may rest for the present, but it is definitely indi- 
cated that new and ample collections should be 
made at all localities from which species were 
described by the older authors, not only Blanco, 
but also others. Interpretation of old species by 
careful study of the florulas of type localities is 
perfectly proper if specimens of the older authors 
have not survived. 

Not only did Georg von Martens look into the 
identity of Blanco's species, he likewise compiled 
an elaborate table showing the known geograph- 
ical distribution in 1866 of all the Algae that had 
been described or reported from tropical Asia 
and the tropical Pacific. This table surely affords 
a key to most of the literature bearing on Philip- 
pine algae that appeared prior to 1866, but also ac- 
counted for the collections of his son Dr. Eduard 
von Martens who accompanied the Prussian 
East Asia Expedition as zoologist. On account 
of his father's special interest in the algae, Eduard 
von Martens collected them whenever he had 
the opportunity. 

Only two Philippine fresh-water species were 
described as new by Georg von Martens, namely 
Cladophora diluta and Cladophora luzoniensis. 
The seaweeds were all considered to have been 
previously known, but there are among them 
first records from various localities. 

A few of the scanty early records of Philippine 
algae were based upon collections of the 
American Exploring Expedition under the com- 
mand of Charles Wilkes. The botanical collec- 
tions of the Wilkes Expedition were made by 
Charles Pickering, general naturalist and ethnobo- 
tanist, author of The Chronological History of 
Plants: Man's Record of his own Existence 
illustrated through their Names, Uses, and Com- 
panionship, William D. Brackenridge, who 
wrote a volume on the ferns, and William Rich, 
who was primarily interested in collecting flower- 
ing plants. Because there was no especialist in the 
lower cryptogams among the naturalists of the 

t Made possible through the Guggenheim fellowship and the University of the Philippines special detail fellowship during 


Presented by G.F. Papcnfuss. 

t Secretary of the Graduate School and Associate Professor in Botany, University of the Philippines. 

1 See J.G. Agardh, 1851, in Species Genera et Ordines Floridearum, 2: 268. 




Expedition (and such were few at that early date), 
the collections of algae were meagre. They were 
determined by Bailey and Harvey 2 , in whose report 
there are two lists, one of the large forms con- 
taining numbers of Philippine reds and browns, 
only one of them new, namely Dictyota dichotoma, 
and a second longer list containing several new 
species of diatoms, namely Amphitetras favosa, 
Campy lodiscus Kutzingii, Lagena WilUamsonii, 
and Triccratiwn orientate. 

All of the diatoms were listed as having come 
from the Sula Sea, without designation of place, 
but are fairly considered as part of the Philippine 
flora, as they were obtained as epiphytes on the 
larger algae. There is to be found in the general 
report of the Expedition, which was written by 
Wilkes himself, a clear indication that shore 
collecting was done at only one Philippine 
locality. That was at Marongas Island, indicated 
on Wilkes* map of Jolo (Sooloo) Island as lying 
across a narrow strait of the northeast coast of 

An English resident of Manila during the 
Spanish regime was Hugh Cuming, who made 
the most extensive collections down to the time 
of the American occupation. His algal collections 
were described by C. Montagne. They were 
widely distributed from Kew, and it was possible 
even as late as the early years of the present cen- 
tury for Dr. E.D. Merrill to secure some of the 
duplicates for the herbarium of the Bureau of 
Science, which had been built up vigorously dur- 
ing the period prior to World War II. The 
Philippine algae of the famous British "Challenger 
Expedition" were enumerated by Dickie in 1876 
and 1877. The specimens were presumably in 
English herbaria. A good many algae were 
accumulated by the Bureau of Science collectors 
and by botanists connected with the University 
of the Philippines. This included Walter Shaw, 
especially interested in the Volvocaceae, who 
wrote on that group and published new Philip- 
pine genera, namely: Campbellosphaeria, Janeto- 
sphaeria, Merrillosphaeria, and Copelandosphaeria. 
It was fortunate for the beginning of phycological 
study that some of the algae of the Bureau of 
Science and University herbaria had been sent out 
as exchanges or on loan prior to World War II 
and are still to be found in various herbaria a- 

During the early part of the American period 
the most extensive collecting of Philippine algae 

was carried out by the Dutch Siboga Expedition. 
There was a general report by Mme. Weber van 
Bosse on all of the collections, and monographs 
of special groups, the Codiaceae by A. and 
E.S. Gepp, the genus Halimeda by E.S. Barton, 
and the remarkably well represented corallines 
by A. Weber van Bosse and M. Foslie. The 
Siboga Expedition dredged very extensively at 
charted stations throughout the Sulu Sea, and 
the Foslie contributions to our knowledge of the 
group are basic. The specimens are presumably 
in Holland and elsewhere in Europe. They were 
never represented in the Philippine herbaria. 

The algae of the voyage of the Italian Vettor 
Pisani were described by A. Picconc in 1886. 
Later in 1889, three more species were added in 
the record of Philippine algae. That ended the 
period of visits of oceanographic expeditions, so 
far as reports on algae were concerned. 

During the incumbency of an exchange profes- 
sorship at the University of the Philippines in 1935, 
H.H. Bartlett began a systematic effort to collect 
the algae. Many localities were visited by him, 
his colleagues, and students. The collections have 
so far been studied only partially, the Chlorophy- 
ceae by Dr. W.J. Gilbert, the genus Galaxaura 
by Dr. Ruth Chou, and various Myxophyceae 
by the present writer. Fortunately, duplicates of 
almost everything went to the University of 
Michigan, and therefore remain as a basis for 
further study. A collection of minute epiphytes 
had been picked off of larger algae and mounted 
on slides. These only were lost, but they can 
doubtless be replaced from the larger herbarium 
specimens. A series of plankton collections were 
also lost. The localities represented have been 
listed by Gilbert in his doctoral dissertation. 
Most of the Myxophyceae were collected by the 
present writer. He was induced by the urgency 
of the nuisance caused in fish ponds by blue- 
green algae to concentrate on that group. Hun- 
dreds of collections were made, which were 
largely studied taxonomically by Drouet, and 
which appeared in numerous papers prepared by 
the writer. But the more extensive is Drouet 
and Daily's "Revision of the coccoid Myxo- 
phyceae" (12) which cited all myxophycean 
unicellular forms so far known from the Philip- 
pines. Thanks to the circumstances of duplicates 
having gone abroad, the vouchers for Philippine 
collecting in this group down to 1941 have not 
been lost. 

2 Summary of the life of J.W. Bailey and W.H. Harvey, each a pioneer in his chosen botanical career at the time, may 
be found respectively in Appleton's Cyclopaedia of American Biography, V. 1, 1888 and Dictionary of National Biography, 
V. 25, 1891. 



Bartlett was again in the Philippines in 1940-41, 
and although engaged in agronomic work, he 
arranged for a pearl diver (Balhani, Moro of 
Siasi) to make bulk collections at various places 
in the Sulu Sea that could be reached by native 
vinta from Zamboanga. The collecting was 
supervised in large part by Professor Jose S. 
Domantay of the Bureau of Fisheries. All of 
these collections were preserved in formaldehyde 
in 55 five-gallon cans. The huge task of making 
them into herbarium specimens was undertaken 
by the University of Michigan under the super- 
vision of Dr. Wm. Randolph Taylor. They have 
not yet been studied, but sets will be returned 
eventually to the Philippines for the University 
and the National Herbarium. 

In spite of the discouraging loss of all of the 
botanical collections and the libraries of the 
Bureau of Science and the University of the 
Philippines during the war for liberation of 
Manila, an immediate effort was made in 1945 
to make a new start. This was greatly aided by a 
grant-in-aid to the present writer from the Ameri- 
can Philosophical Society, which enabled him to 
continue his own researches on the Myxophyceae. 
Several of his students developed a sufficiently 
strong interest so that they were able to publish 
contributions or deposit unpublished master's 
theses, notably J.D. Soriano, M. Cantoria, 
V. Aligaen, E. Medina, M. Valero, and V.G. 
Viola. Needless to say, the publications and 
theses of these immediate post-war workers could 
not have been prepared in the absence of all 
reference materials without the friendly and 
much appreciated collaboration of authorities 
abroad, among whom should be especially 
mentioned Dr. Francis Drouet at the Crypto- 
gamic Herbarium of the Chicago Natural History 
Museum and Dr. Wm. Randolph Taylor of the 
University of Michigan. 

In order to maintain impetus, the Algological 
Society of the Philippines was organized in 
June 1956, with sixteen initial members. Later 
the names were added of several persons who 
were interested in algae as sources of food, in 
their uses in pharmacy and various fertilizer ana- 
lyses in chemistry, or in other economic aspects 
of the group such as their cultivation in fishponds. 
Dean Patrocinio Valenzuela, Executive Secretary 
of the National Research Council of the Philip- 
pines, and Dr. Deogracias Villadolid, former 
Director of the Bureau of Fisheries and presently 
engaged in fish culture, were made honorary 
members to furnish moral support and necessary 
encouragement to the new organization. Among 


the enthusiastic members are Professor Domantay 
who collaborated with Bartlett in organizing and 
supervising the Sula Sea collecting in 1940-41, 
Mr. I. A. Ronquillo of the Bureau of Fisheries, 
Mr. M. Palo of the Institute of Science and 
Technology, Mr. Z.R. Torres of the University of 
the East, and Dr. J.V. Santos of the Department 
of Botany, University of the Philippines. 

As the members of this Society could do little 
at the start, except to collect and to make ecolog- 
ical and economic observations, they have con- 
ceived the idea of building up a "bank" of well 
duplicated numbered specimens, only roughly 
identified as to family or genus, from which 
specialists abroad may receive material for use 
in monographic or regional studies. It is hoped 
that there will be requests from members of the 
Subcommittee on Algae of the Committee on 
Botany of Pacific Science Council for material to 
be used in their various researches. As fast as 
material from this "bank" is identified, remaining 
specimens will be incorporated with the organized 
herbaria of the Philippine National Museum, 
the University of the Philippines, and any other 
institutions that participate in the project. The 
preparation of "Algae Philippinae Exsiccatae" 
is soon to be started with the cooperation of 
Dr. Mason Hale, Associate Curator of the Divi- 
sion of Cryptogams, U.S. National Herbarium 
of the Smithsonian Institution. 

There is every reason why there should be 
immediate cooperation with other countries of 
Southeastern Asia and the Pacific. There is so 
large a widely distributed element in the algal 
floras of the tropical countries that are neighbors 
to the Philippines that an understanding of geo- 
graphical and ecological relationships requires 
the most extended study possible. Furthermore, 
there are various new political entities in the Old- 
World tropics with newly established scientific 
institutions which might like to cooperate closely 
with the Philippines, or to organize regional 
survey work independently in a similar manner. 

In the rebuilding of Philippine references and 
research collections an effort is being made to 
secure for Manila institutions as much duplicate 
prewar material as can be found abroad. The 
fortunate circumstance that much still remains 
for distribution from the University of Michigan 
has already been mentioned. The greatest pos- 
sible cooperation has been realized from Dr. 
Drouet of the Cryptogamic Herbarium, Chicago 
Natural History Museum, who has sent to Manila 
whatever could be spared of duplicates sent him 
prior to the destruction of the Philippine institu- 



tions. Dr. Tiffany of Northwestern University 
turned over to the writer all his accumulated 
articles on the Myxophyceae for library refer- 
ences. There could be no greater proof of the 
value of collecting abundant material and distri- 
buting it widely. What is destroyed in one place is 
preserved in another. As a result of inter-institu- 
tional cooperation between the Philippines and 
the United States, for example, the voucher 
specimens for 183 Philippine records of Myxo- 
phyceae, including types, have been preserved. 


(1 ) Aligaen, V., 1954, Studies on the Algae of 

the Brackish Water Fishponds of Iloilo 
and Vicinity. Master of Science thesis 
(in press), University of the Philippines. 

(2) Bailey, J.W., and Harvey, W.H., 1851, Des- 

cription of Algae from the Wilkes 
United States Exploring Expedition in 
Proceedings of the Meeting for Dec. 4, 
1850, Proc. Boston Soc. Nat. Hist., 3: 

(3) - - , 1862, Uni- 

ted States Exploring Expedition during 
the Years 1838-1 842, under the Command 
of Charles Wilkes, U.S.N., XVII, Phila- 
delphia, pp. 153-182. 

(4) Barton, E.S., 1901, The Genus Halimeda, 

Siboga Expeditie Monographic, 60, 32 
pp., 4 pis. 

(5) Blanco, M., 1837, Flora de Filipinas, Segun 

el Sistema Sexual de Linneo. LXXV11I- 
887 pp., Manila. (There is also an 
1845 edition.) 

(6) Britton, M.E., 1948, New Species of Oedogo- 

nium from Leyte, the Philippine Islands, 
Amer. Jour. Bot., 30 (14): 715-719, 
16 figs. 

(7) Cantoria, M., Valenzuela, P., and Velasquez, 

G.T., 1951, Pharmacopoeial Properties 
of Agar from Three Philippine Seaweeds, 
Jour. Phil. Pharm. Assoc., 38: 187-190, 
3 figs. 

(8) Chou, Ruth C, 1945, Pacific Species of 

Galaxaura, I : Asexual Types. Papers of 

the Mich. A cad. Sci. Arts and Letters, 

30:35-55, 11 pis. 
(9) -- , 1947, Pacific Species of 

Galaxaura, II: Sexual Types, Ibid., 31 

(1945): 3-24, 13 pis. 
(10) Dickie, G., 1876, Contributions to the 

Botany of the Expedition of H.M.S. 

"Challenger" Algae Chiefly Polyne- 
sian, Jour. Linn. Soc. Bot., 15: 235-246. 

(11) __ ._. , 1877, Supplementary 

Notes on Algae Collected by H.N. 
Moseley, M.A. of H. M. S. "Challenger" 
from Various Localities, Jour. Linn. Soc. 
Bot., 15: 486-489. 

(12) Drouet, F., and Daily, W.A., 1956, Revision 

of the Coccoid Myxophyceae, Butler 
Univ. Bot. Stud., 12: 1-218. 

(13) Gepp, A., and Gepp, E.S., 1911, The 

Codiaceae of the Siboga Expedition, 
Siboga Expeditie Monographic, 62, 150 
pp., 22 pis. 

(14) Gilbert, W.J., 1942, Studies on the Marine 

Chlorophyceae of the Philippines. Doc- 
tor of Philosophy thesis (unpublished), 
University of Michigan. 

(15) Martens, G. von, 1866, Die Proussische 

Expedition nach Ost-Asien , Bot. 
Thcil. Die Tange 4 plus 152 pp., 3 pis. 
K. Geheime, Berlin. 

(16) Medina, E., 1955, Studies on the Siphonales 

of Puerto Galera, Oriental Mindoro. 
Master of Science thesis (in press), 
University of the Philippines. 

(17) Merrill, E.D., 1918, Species Blancoanae, a 

Critical Revision of the Philippine 
Species of Plants Described by Blanco 
and Llanos, Dept. Agr. and Nat. Res., 
Bu. Sci., Manila, 12, 423 pp., 1 map. 

(18) Piccone, A., 1886, Alghe del Viaggio di 

Circumnavigazione della Vettor Pisani, 
Genoa, 97 pp., 1 map. 

(19) , 1889, Nuove Alghe del 

Viaggio di Circumnavigazione della Vet- 
tor Pisani, Roma, R. Ace. Lined Mem. 
Series 4, 6: 10-63. 

(20) Rabanal, H.H., 1949, The Culture of Lab- 

lab, the Natural Food of the Milkfish Fry 
and Fingerlings under Cultivation, The 
Technical Bulletin, 18, Bu. of Printing, 

(21) --, and Montalban, H.R., 1953, 

The Growing of Algae or "Lumut" in 
Bangos Fishponds, Philippine Fisheries, 
pp. 142-152, 3 figs. 

(22) Shaw, W.R., 1919-1922, Papers on Volvo- 
caceae of the Philippines, four new 
genera in separate articles, Phil. Jour. 
Sci., respectively: Cawpbellosphaeria, 
15 (6): 493-520, 2 pis., 1 fig.; Janetos- 
phaeria, 20 (5): 477-508, 5 pis., 5 figs.; 



Merrillosphaeria, 21 (1): 87-129, 8 pis., 
1 fig.; and Copelandosphaeria, 21 (2): 
207-232, 4 pis., 2 figs. 

(23) Soriano, J.D., 1953, Myxophyceae of Panay 

and Ncgros Islands, Nat. and A pp. Set. 
Bull., 13(1 and 2): 3-57, 3 pis. 

(24) , and Velasquez, G.T., 1952, 
Studies on the Myxophyceae of Manila 
and Vicinity, Ibid., 12 (1): 1-93, 4 pis. 

(25) Tiffany, L.H., 1951, Two New Oedogonia 

from the Philippines, Chicago Acad. 
Sci., 82: 81-83, 1 pi. 

(26 ) Valero, M., 1956, Preliminary Studies on 

the Algae of U.P. Site, Quezon City. 
Master of Science thesis (unpublished). 
University of the Philippines. 

(27) Velasquez, G.T., 1940-41, Papers on Fila- 

mentous Myxophyceae of the Philip- 
pines 1, II, and III, Nat. and App. Sci. 
Bull., respectively: 7 (3): 269-271 : 8 (2): 
189-200: and 8 (3): 203-210. 



, 1948, Survey of the Algae 
and Economic Algal Resources of the 
Philippines, The Amcr. Philosophical 
Society Year Book, pp. 154-155. 

, 1950, Studies on the Myx- 
ophyceae of Puerto Galera and Vicinity, 
Nat. und App. Sci. Bull., 10(4): 309-328. 

, 1952, Algal Pollutions 
from the Ponds of Puerto Galera, Orien- 
tal Mindoro, Ibid., 12 (3): 239-251. 

, 1952, 
of the Philippines, 

Seaweed Resources 
Proc. of the First 

International Seaweed Symposium, In- 
stitute of Seaweed Research, Scotland, 
pp. 100-101. 

(32) , 1953, Studies on the 
Marine Algae of the Philippines. Ab- 
stract, Proceedings of the Eighth Pac. 
Sci. Cong., pp. 205-206. 

(33) Villadolid, D.V., and Villaluz, O.K., 1953, 

A Preliminary Study on Bangos Cultiva- 
tion and Its Relation to Algal Culture in 
the Philippines, Bu. of Printing, Manila, 
Pop. Bull., 30, pp. 3-16. 

(34) Viola, V.G., 1956, The Chlorophyceae of 

Puerto Galera and Vicinity, Their Dis- 
tribution and Reproduction. Master of 
Science thesis, University of the Philip- 
pines. (With permission to be read in 
the Ninth Pacific Science Congress, 

(35) Weber van Bosse, Ann., 1913-28, Listc des 

algues du Siboga, I: Myxophyceae, 

Chlorophyceae, Phaeophyceae , pp. 

1-1 86, pis. 1-5(1913). II:Rhodophyceae, 
premiere partie, Protoflorideae, Nema- 
leonales, Cryptonemiales, pp. 187-310, 
pis. 6-8 (1921); secondc partie, Cerami- 
ales, pp. 311-392, pis. 9-10 (1923); troisi- 
cmc partie, Gigartinales et Rhodymeni- 
ales , pp. 393-533, pis. 11-16 (1928), 
Siboga Expeditie Monog., 59, E.J. Brill, 

(36) , and Foslie, M., 

1900, The Corallinaceae of the Siboga 
Expedition, Sihoga Expeditie Mono- 
graphie, 61, 110 pp., 16 pis., 34 text figs. 


G.F. PAPENFUSS: Perhaps Dr. Santos can tell us what 
has happened to Blanco's material. 

j.v. SANTOS: 
the war. 

It was destroyed by fire in Manila during 

I should explain that the algal society mentioned by 
Dr. Velasquez has been formed from members from 
zoology, medical, and fisheries departments. The algal 
collection is increasing from time to time, and Dr. 
Velasquez is organizing the identifications. 






University of British Columbia, Vancouver, Canada. 

Although Schmitz (14) and Schmitz and Fal- 
kenberg (15), as well as earlier authors, had sub- 
divided the family Rhodomelaceae into a number 
of subfamilies, Falkenberg's (2) monograph on 
the Rhodomelaceae was the first noteworthy 
attempt at a broad phylogenetic study of this 
largest family of Rhodophycophyta in which 
there are some 1,200 species now recognized. 
Oltmanns (8, 9), Rosenberg (11), Kylin ?tf, 7), 
Fritsch (3), and Scagel (12) have attempted to 
bring the group in line with more modern con- 
cepts of classification in the Florideophycidae, 
but each, with some modification, has essentially 
followed Falkenberg's division of the family into 
subfamilies. Rosenberg (II) removed the Dasy- 
aceae from the Rhodomelaceae and established 
this group as an autonomous family, the Dasy- 
aceae, and Scagel (12) has attempted, on the basis 
of a study of certain of the dorsiventral Rhodo- 
melaceae, to further assess phylogenetic relation- 
ships in the family. Kylin (7 ) has departed to the 
greatest extent from the system of classification 
proposed by Falkenberg (2). It has been possible 
by further study of some of the genera in this 
large group to place a number of entities formerly 
of uncertain position. However, there still exists 
in this family a broad field for research on a 
generic level before a satisfactory approach can 
be made to the solution of the many specific 
problems and before the merits of the phylogene- 
tic relationships can be adequately weighed. As 
Oltmanns (8, 9) has stressed, one must have a 
detailed knowledge of the family as a whole before 
a reassessment of such a large group can be made. 
Many of the genera, even as they have now been 
rearranged or subdivided by Kylin (7), still in- 
clude obscure and little-known plants which 
have never been adequately studied. There seems 
little doubt that further rearrangements and sub- 
divisions of this large family will be warranted, 
but until more of the entities have been studied 
in greater detail and in all stages of development 
there seems little justification in proceeding fur- 
ther at this time in an attempt to regroup genera 
into subfamilies. 

The position of the Rhodomelaceae in the 
classification system, as for the majority of the 
groups in the Florideophycidae, is based essen- 

tially on embryological characters proposed by 
Schmit/ (13). More recent studies, particularly 
on the female reproductive apparatus, have 
further emphasized the importance of using 
these structures for the interpretation of phy- 
logenetic relationships on both ordinal and family 
levels. The studies of Kylin (4,5 ) and Pa pen fuss 
(10) have particularly stressed the importance 
of the use of the female reproductive system in 
the delimitation and interrelationships of the 
families of algae now included in the Order 
Ceramialcs. It is apparent from these studies 
that the Ceramiales is a highly evolved group. 
In the members of this order the carpogonial 
branch, auxiliary cell, supporting cell, and often 
the central cell, the sterile cells, and in some 
instances even the inner cells of the pcricarpic 
layer, all contribute in varying degree to the 
nutrition of the carposporophyte. Of the four 
families in this order, however, this phylogenetic 
trend in the nourishment and protection of the 
developing carposporophyte is most clearly 
evident in the Rhodomelaceae. In this family, 
the direct fertilization of an auxiliary cell, which 
is formed only after fertilization of the carpog- 
onium, and the early formation and high degree 
of development of the pericarp are well- 
established features. With respect to both re- 
productive and vegetative structures, not only 
the Order Ceramiales, but also the Family Rhodo- 
melaceae is the most clearly defined and most 
highly developed of all groups of Florideophy- 

Chief among the distinguishing characteristics 
of the life-histories of the Rhodomelaceae, as well 
as for the majority of the Florideophycidae, is 
the succession of somatic phases, different from 
that of most other algal groups. This is due to 
the development of the carposporophyte, fol- 
lowing the fertilization of the carpogonium. 
The gametophyte, which bears the sex organs, 
is haploid, and the fertilized carpogonium gives 
rise indirectly to a carposporophyte with diploid 
sporogenous tissue. No reduction division ac- 
companies the production of the carpospores, 
which germinate to give rise to the diploid tetra- 
sporophyte. This tetrasporophyte bears the 
tetrasporangia in which reduction division occurs 



and the haploid tetraspores give rise to male and 
female gametophytes. Thus there are four soma- 
tic phases, two haploid and two diploid. The 
carposporophyte, which is almost like a parasitic 
generation, is much reduced in vegetative develop- 
ment and remains attached to the female game- 
tophyte. Although the tetrasporophyte and the 
male and female gametophytes are quite separate 
phases in the life history, they cannot generally 
be identified morphologically as such, or even 
anatomically until the reproductive cells are 
formed. For this reason, there is an even greater 
opportunity in such a group for studying vege- 
tative characteristics both in the haploid and 
diploid phases. 

In spite of the large size of the family Rhodo- 
melaceae, there is probably no group of algae 
which shows such a remarkable degree of 
uniformity in the female reproductive organs 
and in the post-fertilization development. Al- 
though the uniformity of the female reproductive 
system is an outstanding feature of the Rhodo- 
melaceae, on the other hand the family shows a 
diversity in vegetative organization not met 
elsewhere in a family of multicellular algae. The 
strictly apical development of the thallus can be 
traced in the young plant by means of the primary 
pit-connections between cell lineages, although 
in older stages secondary pit-connections may 
be formed between adjacent pericentral cell 
derivatives and result in a pseudoparenchymatous 
thallus whose development and organization may 
soon be obscured. Although the development of 
the mature thallus may be completely evident 
from a study of its apices, there are instances, as 
in Placophora, in which the evanescent juvenile 
stages have been found to exhibit an organization 
quite different from that of the mature thallus. 
For this reason it is instructive to study not only 
the mature plants but also the juvenile plants and 
their development before completing a phylo- 
genetic arrangement of genera in this group. 
Although the range in construction of the thalli 
varies from comparatively simple, radially sym- 
metrical plants to elaborate and complex types 
with bilateral symmetry or dorsiventral organiza- 
tion, through a study of the apex of the mature 
plants the polysiphonous theme on which these 
variations are superimposed is readily ascertained 
even in the most highly modified types. 

The family can be subdivided into a number 
of subfamilies which can be arranged in several 
lines of evolution based primarily on symmetry. 
Accumulated evidence supports the theory that 
the subfamily Polysiphonieae is the basic group 
from which all other subfamilies have been 


derived. The ancestral type was probably a sim- 
ple, radially symmetrical, Polysiphonia-like alga 
with five pericentral cells in the polysiphonous 
segments. On this basis the evolutionary lines 
are : the Potysiphonia-series, the basic group from 
which all Rhodomclaccae are probably derived, 
the Lophothalia-, Chondria-, Pterosiphonia-, Her- 
posiphonia-, and Amansia-series. The Polysipho- 
nia-, Lophothalia-, and Chondria-series include 
chiefly radial forms, whereas the Pterosiphonia- 
series includes primarily bilateral forms. The 
Herposiphonia- and Amans la-series include pri- 
marily dorsiventral forms. It is a part of the 
Herposiphonia-series which I shall use particularly 
to illustrate the emphasis which may be placed 
on vegetative characteristics in the group. 

According to Falkenberg (2) this series com- 
prises two subfamilies the Herposiphonieae and 
the Polyzonieae. However, Kyi in (7 ' ) has further 
split up the Herposiphonieae into three groups 
which may be referred to as the Herposiphonieae, 
Placophoreae, and the Streblocladieae. The 
genera included in this series are chiefly dorsi- 
ventrally organized. In some, as in Metamorphe 
(Herposiphonieae), the axes are simple, more or 
less erect, and Polysiphonia-Yike with dorsiven- 
trality very slightly developed. In more advanced 
genera, as Placophora and Amplisiphonia (Placo- 
phoreae), dorsiventrality is very pronounced and 
the mature thalli, which are almost entirely 
prostrate, show extensive congenital coalescence 
of the axes. The Polyzonieae includes genera 
which show strongly pronounced dorsiventrality. 
Many of the latter have acquired the form of 
jungermanniaceous liverworts as the result of 
various types of wing-development (Leveillea), 
the coalescence of determinate laterals (species 
of the "cuneifoliate" group of Dasyclonium 
(=Euzoniella) or a combination of both. 

The initial stages of the prostrate phase of many 
of these dorsiventral forms arise from the basal 
part of an erect, radially or bilaterally symmetri- 
cal, determinate and evanescent juvenile or pri- 
mary axis. But even where there is pronounced 
dorsiventrality, as in Placophora, when repro- 
ductive branches are produced, the latter may 
revert to an erect radial habit. If we interpret 
these juvenile stages as evidence of ontogeny 
repeating phylogeny, then we have additional 
features supporting the theory that the erect, 
radial, Polysiphonia-type represents the prototype 
of the group. One might postulate that in time 
the erect form lost its ability to produce exogenous 
indeterminate laterals. The production of a pros- 
trate indeterminate lateral from the lower part of 
the juvenile axis consequently has survival value, 



and the plant is able to produce an extensive 
thallus despite the fact that it may have lost the 
capacity of producing exogenous indeterminate 
laterals at the apex of an erect primary axis. As 
a result of the development of holdfasts and a 
prostrate habit, this successful, basal, indeter- 
minate branch has gained a still better chance 
of survival. 

To illustrate how one can carry the analysis of 
the vegetative characteristics to the specific level 
I should like to consider in somewhat greater 
detail the genus Dasyclonium (= Euzoniella). 
As far as is known, the dorsiventral phase of the 
thallus of the species of this genus is initiated by 
a primary lateral which arises from the basal 
part of a diminutive, comparatively evanescent, 
radially or bilaterally symmetrical primary axis. 
The two groups within this genus, which have 
been designated (12) as the "incisate" and "cunei- 
foliate" groups, are characterized primarily on 
the basis of the structure of the determinate 
branches. The "incisate" group includes those 
species in which the secondary and subsequent 
orders of branches formed by the determinate 
axes are more or less free from one another at 
maturity. The "cuneifoliate" group includes 
those species in which the secondary and higher 
orders of branches formed on the determinate 
axes are congenitally fused to varying degrees 
at maturity. Kylin (7) has chosen to split up the 
group of species formerly included in the genus 
Euzoniella Falk. 1901 (in the broad sense as the 
name was proposed for conservation as against 
Dasyclonium J.Ag.,7; by Silva,/6,/7) into two 
groups (Dasyclonium J.Ag., sensu Kylin, 7 and 
Euzonia Kylin, 7). These two groups are those 
which have already been designated as the 
"incisate" and "cuneifoliate" groups. I do not 
believe that the separation of this group of species 
into two genera is justified. Further confusion 
results in properly assigning certain entities, one 
of which Kylin apparently overlooked, [Dasyclo- 
nium palmatifidum (Grun.) n. comb. (= Euzoniella 
palmatifida (Grun.) Cuoghi-Cost.), D. harveya- 
num (Decne ex Harv.) Kylin (= Euzoniella har- 
veyana (Decne ex Harv.) Falk.) and D. ocellatum 
(Yendo) n. comb. (= Euzonia ocellata (Yendo) 
Kylin) ] to Dasyclonium sensu Kylin and his new 
genus Euzonia. Although a good case may be 
made for conserving Euzoniella as proposed by 
Silva (16), as this proposal was subsequently 
rejected in favour of Dasyclonium, it would now 
seem best to reject Kylin's division of the group 
and refer all species to the one genus Dasyclonium 
J.Ag. (= Euzoniella Falk.). On the basis of the 
structure and coalescence of the determinate 

branches, the species of this genus can be arranged 
in an evolutionary series whose equal for com- 
pleteness is probably not encountered elsewhere 
in the family ( 12). Any attempt to separate these 
two groups serves only to obscure their natural 
relationships. It should be noted that in the 
"incisate" group, which Kylin (7) refers to Dasy- 
clonium, he has included D. harveyanum (Decne 
ex Harv.) Kylin. However, the lowermost 
lateral of the determinate branch of this species 
is further branched, a feature which is character- 
istic of the species of the "cuneifoliate" group 
and, although Dasyclonium harveyanum might 
be regarded as a transitional species, it should 
more properly have been placed by Kylin in the 
genus Euzonia. On the other hand he has placed 
Dasyclonium ocellatum (Yendo) n. comb. (= Euzo- 
niella ocellata Yendo ; = Euzonia ocellata (Yendo) 
Kylin), in which there is no coalescence of mono- 
siphonous laterals, with the "cuneifoliate" group, 
whereas this species is most closely related to 
D. incisum (J.Ag.) Kylin (= Euzoniella incisa 
(J.Ag.) Falk.). Furthermore, Kylin has omitted 
any mention of D. palmatifidum (Grun.) //. comb. 
which also forms a natural link between the 
"incisate" and "cuneifoliate" species. Dasyclo- 
nium palmatifidum has only incipient stages of 
coalescence in the laterals on the determinate 
branch, but it has the branching habit of the 
lowermost lateral of the determinate branch 
typical of the "cuneifoliate" group. Thus it can 
be shown that there are species of the genus 
Dasyclonium which illustrate all stages in transi- 
tion from the monosiphonous, uncoalesced con- 
dition of secondary laterals through to a poly- 
siphonous (or partially polysiphonous) and 
completely coalesced condition. The develop- 
ment of the spermatangia axes in Dasyclonium 
incisum, D. palmatifidum, and D. cuneifolium 
(Mont.) n. comb. (= Euzonia cuneifolia (Mont.) 
Kylin; = Euzoniella cuneifolia (Mont.) Falk.) are 
also identical, as are the number and arrangement 
of cover cells formed during the development 
of the tetrasporangium. All of these features 
further support the argument that these entities 
are all species of the same genus. 

Dasyclonium incisum apparently is one of the 
most primitive species in the genus, and may 
have served as the prototype from which D. 
flaccidum (Harv.) Kylin and D. ocellatum have 
been derived by reduction of the polysiphonous 
laterals on the determinate branches to the mono- 
siphonous unbranched condition. On the other 
hand, it seems likely that a monosiphonous con- 
dition throughout may represent the prototype 
for the genus as a whole. Dasyclonium bipartitum 



(Hook. f. et Harv.) Kylin [= Euzoniella bi- 
partita (Hook. f. et Harv.) Falk.] may also be 
considered as having been derived through re- 
duction from D. incisum. In this instance it is 
the number of secondary laterals that has been 
reduced rather than the polysiphonous character 
of the laterals that has been lost. In D. biparti- 
tum only the two lowermost secondary laterals 
are initiated. 

The next probable step in the evolution of the 
determinate branch is indicated in Dasyclonium 
harveyanum. In this species several orders of 
branches, which remain free from one another at 
maturity, are formed from the lowermost lateral 
of the determinate appendage. This branching 
characteristic is also found throughout the 
"cuneifoliate" group and suggests that D. 
harveyanum forms a connecting link, as has 
already been suggested, between the "incisate" 
and "cuneifoliate" groups. In the "cuneifoliate" 
group, various degrees of congenital coalescence 
of the secondary laterals have occurred, resulting 
in the evolution of a monostromatic (for the 
most part) determinate appendage. The incipient 
stages of this coalescence and monostromatic 
character are indicated in D. palmatifidum. This 
congenital coalescence of the laterals is more 
evident in D.flabellifera (J.Ag.) n. comb. [-Euzo- 
niaflabellifera (J.Ag.) Kylin; = Euzoniella flabel- 
lifcra (J.Ag.) Laing] and is even more pronounced 
in D. ovalifolium (Hook. f. et Harv.) n. comb. 
[=Euzonia ovalifolia (Hook. f. et Harv.) Kylin; 
^Euzoniella ovalifolia (Hook. f. et Harv.) Falk.], 
D. cuneifolium and D. adiantiformis (Decne) n. 
comb. [-Euzonia adiantiformis (Decne) Kylin; 
^Euzoniella adiantiformis (Decne) Falk.]. 

In summary one can say that the female re- 
productive structures, which are strikingly uniform 
throughout the family Rhodomelaceae, appear 
to be of least phylogenetic significance at the 
subfamily, generic, and specific levels. The 
diversity in vegetative construction that is asso- 
ciated with this uniformity in the structure of the 
reproductive organs in the Rhodomelaceae, as 
well as in the Ceramiales as an order, indicate 
that the group is a long-established one, whose 
representatives have undergone pronounced 
evolutionary development in the vegetative system 
in comparatively recent times. Of the families 
constituting this order, the Rhodomelaceae is 
the most highly evolved. 

Symmetry relationship is the most fundamental 
feature on which a separation of the subfamilies 
into the various probable lines of evolution can 
be made. Bilaterality and dorsiventrality appear 


to have been achieved more than once within the 
family along entirely different lines of evolution, 
but in general there appears to have been an 
advance from simple forms with a radial organ- 
ization as in the Polysiphonieae to bilateral types 
as in the Pterosiphonieae, and finally to elaborate 
forms with dorsiventral organization as in the 

The method of formation, type, pattern of 
distribution, and the degree of congenital fusion 
or subdivision of the laterals are of secondary 
importance, but are significant at the generic and 
specific levels in distinguishing taxa. 

The genus Euzonia Kylin is rejected because 
the species included in it form only a part of an 
evolutionary series of closely related entities 
hitherto referred to the genus Euzoniella Falk. 
Furthermore, because of the additional confusion 
that has resulted by referring only a part of this 
same series to Dasyclonium J.Ag. (sensu Kylin), 
and because an earlier proposal to conserve Euzo- 
niella has been rejected, it would seem best to 
refer all entities to the one genus Dasyclonium 


(1) Agardh, J.G., 1894, Analecta Algologica 

Contin. IL Lunch Univ. Arsskr. 30 (Afd. 
2, Nr 7), 98 + 1 pp., 1 pi. 

(2) Falkenberg, P., 1901, Die Rhodomelaceen 

des Golfes von Neapel und Angrenzen- 
den Meeresabschnitte, Fauna und Flora 
des Golfes von Neapel, 26: i-xvi, 1-754, 
10 figs., 24 pis. 

(3) Fritsch,F.E., 1945, The Structure and Repro- 

duction of the Algae, Vol. 2, Cambridge, 
xvi -f 939 pp., 336 figs. 

(4) Kylin, H., 1928, Entwicklungsgeschichtliche 

Florideen Studien., Lunds Univ. Arsskr. 
N.F. 24 (Avd. 2, Nr 4), 127 pp., 64 figs. 

(5) _ , 1930, Uber die Entwicklungsges- 

chichte der Florideen, Lunds Univ. 
Arsskr. N.F. 26 (Avd. 2, Nr 6), 104 pp., 
56 figs. 

(6) _., 1937, Anatomic der Rhodophy- 

ceae. In Linsbauer, K., Handbuch der 
Pflanzenanatomie, II. Abt. Band VI, 
2 Teilband, Berlin, viii -f 347 pp., 252 

(7) , 1956, Die Gattungen der Rhodo- 

phyceen, xv + 673 pp., 458 figs, (in- 
cluding Nachtrag by E. Kylin), Gleerup. 



(8) Oltmanns, F., 1904, Morphologic und Bio- 
logic der Algcn, Ed. 1, Spezieller Teil. 
Jena, vi + 733 pp., 476 figs. 

(9) , 1922, Morphologic und Bio- 
logic der Algen, Bd. 2, 2nd ed. Jena, 
iv + 439 pp., 325 figs. 

(10) Papenfuss, G.F., 1944, Structure and Taxo- 

nomy of Taenioma\ including a discus- 
sion on the Phylogeny of the Ceramiales. 
Madrona, 7: 193-214, inch pis. 23 and 
24, 1 fig. 

(11) Rosenberg, T., 1933, Studien iiber Rhodo- 

melaceen und Dasyacecn, Lund (Thesis), 
87 pp., 25 figs. 

(12) Scagel, R.F., 1953, A Morphological Study 

of Some Dorsiventral Rhodomelaceae, 
Univ. Calif. Publ Bot., 27 (1): 1-108, 
20 figs. 

(13) Schmitz, F., 1883, Untersuchungen iiber die 

Befruchtung der Florideen, Sitzungsber. 
K. Preuss. Akad. Wiss. Berlin, 1883: 
215-258, pi. 5. 

(14) , 1889, Systematische Ubersicht 

der Bisher Bekannten Gattungen der 
Florideen, Flora, 72: 433-456, pi. 21. 

(15) , and Falkenberg, P., 1897, Rho- 
domelaceae. In Engler, A., and Prantl, 
K., Die Natiirlichen Pflanzenfamilien.... 
Teil l,Abt.2: 421-480, figs. 240-266. 

(16) Silva, P.C, 1950, Generic Names of Algae 

Proposed for Conservation, Hydrobio- 
logia, 2:252-280. 

(17) , 1953, A Review of Nomenclatu- 

ral Conservation in the Algae from the 
Point of View of the Type Method, Univ. 
Calif. Publ. Bot., 25 (4): 241-324. 



Symposium: Ethnobotany of Thailand and Contiguous Countries 


Institute for Regional Exploration, Ann Arbor, Michigan, U.S.A. 

The subject matter of the current symposium 
is ethnobotanical. It seems, however, that this is 
also the subject matter of at least several other 
sessions of the Congress. I have counted two 
dozen papers given elsewhere which are directly 
concerned with ethnobotany, including such 
topics as the origin of paddy field culture in 
ancient Japan, the study of variation of taros and 
kumaras and its possible ethnobotanical signi- 
ficance, a symposium on vernacular names (Bota- 
ny Division), several papers on the geographical 
differentiation and origin of barley, banana trees, 
and other cultivated plants (Agriculture) as well 
as papers forming a part of the symposium on 
the humid tropics, sessions on coconut problems 
and on shifting agriculture. 

The dispersion of ethnobotanical reports 
throughout the Congress is unavoidable, of course, 
for it is probably the only way of including 
ethnobotanical data in specialized symposia. In 
other respects it is unfortunate. It means that 
many papers will not reach certain persons who 
are interested in them until after they are pub- 
lished, which may be a matter of years. Further- 
more I have noticed in attending sessions 
that many discussions suffer because they lack 
authoritative opinions which probably would be 
present if there were a concentration of ethno- 
botanical material in a single, jointly-sponsored 
session or sessions. 

Today, I would like to begin our symposium 
on the ethnobotany of Thailand and contiguous 
countries by offering a definition of our subject. 
It is evident, I think, that there is a difference of 
opinion concerning the nature and scope of 
ethnobotany as witnessed, for example, by the 
diversity of subjects covered in our symposium 
this morning. For purposes of discussion, I 
propose a definition of ethnobotany that is largely 
accepted in America, that is: ethnobotany is a 
study of the interrelations of primitive man and 

There is, I believe, a distinction between eth- 
nobotany and economic botany. Ethnobotany 
should deal with verbal traditions only, except 
for those traditions which have recently found 

their way into the literature, whereas economic 
botany is more concerned with modern concepts 
of the science of plants and their uses. The 
word "primitive" often carries with it unfortunate 
social implications; it has different meanings for 
different people. Yet it is nevertheless our best 
key to the real content of ethnobotanical study. 
Ethnobotany is primarily concerned with the 
interrelations of plants and primitive man in the 
sense that "primitive" denotes only a lack of any 
written language and therefore the preservation 
of traditions by verbal means alone. This is, 
I think, the accepted anthropologicall view. 

Chemical analysis of useful plants is not usually 
considered a part of ethnobotanical work, where- 
as the following are: the study of plant debris in 
archaeological deposits, studies of plant patterns 
in the vicinity of old village sites, pollen analysis, 
and radiocarbon dating. 

The earliest use of the term "ethnobotany" was 
by J. W. Harshberger in 1895. It was first used 
narrowly in reference to the use of plants by 
aborigines. More recently, most authors agree 
that ethnobotany should deal "not only with 
plant uses, but with the entire range of relations 
between primitive man and plants." (V. H. Jones, 
1941, p. 219). 

It is evident, therefore, that our study must be 
a broad one, for man and plants are co-existent 
over a broad spectrum. There is necessarily 
ecological interaction between them, as Jones 
points out. Perhaps of even greater significance 
is the fact that there are cultural forces that must 
be reckoned with, and these make our study 
uniquely different from animal ecology. It is 
necessary that we make use not only of bio-ecolo- 
gical concepts in dealing with plants and man in 
any ethnobotanical work, but also we must bring 
into our study the various concepts which an- 
thropologists have regarding cultural dynamics. 

It goes without saying that ethnobotany is an 
interdisciplinary study that overlaps a number of 
established scientific fields. Perhaps, as some 
have suggested, the real function of ethnobotanic- 
al work is to iilucidate the interaction of primi- 
tive man and plants on a cultural level, not on a 



biological level, and we should leave the biologi- 
cal interpretations to the plant and animal scien- 
tists. To my mind, however, such a restriction 
on the scope of ethnobotany is a step in the 
wrong direction. Ethnobotanical study will be 
of far greater significance if it seeks to overlap, 
if it brings together various approaches of various 
disciplines for the better understanding of both 
the cultural and the biological factors involved 
in the interrelations of plants and primitive man. 

The main point I would like to leave from this 
perhaps overly-long dwelling on already pub- 
lished ideas is that ethnobotany is really an inte- 
gral part of the newly developing science which 
various authors have called "human ecology" 

and "cultural ecology." The content, philoso- 
phies, and methods of this new science are 
yet to be satisfactorily defined. Various authors 
have their own views of just what "human 
ecology" is. But from the strides which bio- 
ecology has made toward illucidating for us the 
forces of Nature, and their effects upon biota 
and environment, I feel certain that the study we 
are beginning to call human ecology will be able 
to do the same for a better understanding of the 
forces of culture. It is significant that the term 
"ecology" implies that specialists will be involved 
in the study not only from anthropology but from 
the biological sciences as well. Ethnobotanists 
will undoubtedly be called upon to make major 
contributions to the combined effort. 





University of Michigan, Ann Arbor, Michigan, U.S.A. 


"The Nature and Status of Ethnobotany" was 
published by Professor Jones in 1941, when it 
appeared in Chronica Botanica, Volume VI, 
Number 10, pp. 219-221. Because it contains 
some pertinent thoughts on the content and status 
of ethnobotany which arc worth repeating for 
the benefit of this symposium on "The Ethno- 
botany of Thailand and Contiguous Countries," 
I have taken the liberty, with Professor Jones' 
permission, of extracting portions for this paper. 

Jones points out in his original article that a 
large amount of data associated in some manner 
with the relationships of primitive man and plants 
has accumulated in the various literatures of such 
diverse disciplines as botany, anthropology, 
linguistics, agriculture, horticultural science, and 
geography. These data are concerned with econo- 
mic botany, plant lore, properties and value of 
economic plants, the origins of cultivated plants, 
plant remains in archaeological sites, and plant 
names and plant knowledge of primitive peoples. 
In America these various aspects of the study of 
the relationship of aboriginal peoples and plants 
have come to be known under the term "eth- 

Among the students of ethnobotany, the plant 
scientist is primarily interested in knowing what 
plants were and are used by primitive man, how 
they are gathered and utilized, and what effect 
man has had upon the dispersal of plants. The 
former distributions of plants as indicated by 
archaeological deposits should offer the botanist 
valuable data to supplement his own research in 
plant geography. The lists of plant names and 
ethnological and archaeological occurrences of 
plants have been shown to be of value in the study 
of the origin and dispersal of cultivated plants. 
The plant ecologist is interested in the influence 
of primitive man on the plant environment, 
especially the effect of man's activities in dis- 
turbing the otherwise normal processes of plant 
succession. The anthropologist, on the other 
hand, is more interested in the manner in which 
primitive man adapts himself to his plant en- 
vironment, what plants he uses, and how his 
economy, activities, and thoughts are influenced 

t Presented in abstract by T.P. Bank II. 

by the plant world. The cultural implications, 
in other words, are paramount over the botanical. 
The ethnobotanist, speaking of the professional 
worker who specializes in ethnobotany, is most 
useful perhaps in correlating the data on these 
and similar problems and in presenting his results 
in a form that is useful to either the plant scientist, 
the anthropologist, or both. 

Few individuals, either in the United States or 
elsewhere, have given their entire attention to 
ethnobotany alone. Ethnobotany for its own 
sake is practiced by perhaps a handful of workers. 
On the other hand, anthropologists have gone 
into ethnobotany to solve certain problems, and 
botanists have become part-time anthropologists 
for the same reason. Recently there has developed 
in America a healthy and productive cooperation 
among botanists, anthropologists, and linguists, 
and a more concerted attack has been made 
upon ethnobotanical problems of mutual interest. 

Although many ethnobotanical observations 
were made much earlier, it is only since about 
1850 that any great amount of substantial pro- 
gress has been made in the correlation of ethno- 
botanical information. In Europe the most 
notable early work was that of such men as 
Alphonse de Candolle, Ungcr, Targioni-Tozzetti, 
Bretschneider, and Wittmack in applying ethno- 
botanical data to the solution of problems of the 
origins and distributions of cultivated plants. 
More recently, emphasis in Europe has been on 
the development and application of techniques 
for the removal and study of archaeological plant 
materials. Some of the early work in America 
was done by Europeans; but since about 1875, 
American ethnobotany has progressed more or 
less independently. A rather extensive and valu- 
able literature bearing on the relations of the 
Indians of North America and plants has accu- 
mulated. American archaeological plant material 
has yet to receive the attention which it deserves. 
Although little has been done on the ethnobotany 
of South America except in Peru, there would 
seem to be remarkable opportunities in that 


The results of the intensive surveys of the world. In the collecting of such products and 

expeditions of the Soviet Union under the particularly in the obtaining and recording of 

direction of Vavilov seem to indicate that many the knowledge of the natives concerning uses, 

kinds and varieties of cultivated plants distinct properties, cultural treatment, and other such 

in their genetic constitution, adaptations, and information, ethnobotanical experience and the 

virtues from any now in the agriculture of civilized ethnobotanical approach should be valuable, 
man may yet be obtained in various parts of the 





Chief, Section of Botany and Zoology, Forest Products Research Division, Royal Forest Department, Bangkok, Thailand. 

In rural parts of Thailand, thatches are still 
in use, as they are cheap and easy to procure. 
The materials used vary in different parts, but 
fall roughly into three categories: 


In northern Thailand, people collect mature 
leaves of young trees of Mai Pluang (Dipterocar- 
pus tuber culatus ) , abundant in the dry deciduous 
forests. In thatching, these leaves are arranged 
in an overlapping row on a thin piece of bamboo, 
about one metre long, fastened together with 
bamboo or rattan. This kind of thatch, when 
properly seasoned, is not susceptible to insect 
attack and is not inflammable. The thatch lasts 
for two to three years. 


At least four varieties of these leaves are used 
in thatching: 

Kaw (Livistona sp.) is a tall, erect palm of 15 to 
20 metres in height, common in evergreen forest. 
The leaves are fan-shaped and are much used 
by hill tribes who fold them in half into a 
roughly triangular shape which are then ar- 

ranged to overlap each other, being secured to 
the building framework with rattan. The 
thatch does not last long, but as these hill tribes 
move every three or four years, this does not 

Wai (Daemonorops sp.) is a kind of rattan, 
known in certain places as Wai Chak. There 
are two or three species of this rattan whose 
leaves are feather-like in shape with thin spines 
growing on the pinnae and larger spines grow- 
ing on the petioles. The leaves are folded in half 
and placed in pairs to three long rattan canes 
to which they are secured by split rattan bind- 
ing at intervals of 5 cm as illustrated below. 
When a required length has been made, the 
whole sheet is rolled up, ready for laying upon 
the building framework to which it is secured 
by split rattan binding. These thatches last 
from two to three years. 

Chak (Nipa fruticans), a feather-leaved palm 
with underground stem, grows gregariously in 
tidal forests and provides material for roofing 
for people living in the central plain and coastal 
area. The pinnae of matured leaves are cut 
from the axis, one-third of their length is then 




folded and placed overlapping each other on a 
bamboo split and bound with Wai nam (Flagel- 
laria indica), a riperian species of scandent 

Sakhu (Metroxylon sp.), known commercially 
as sago palm, is another form of thatching pre- 
pared in a like manner to Chak. This is com- 
monly used in the Malay peninsular. 


Two kinds are used, Kha (Imperata cylindri- 
cata), commonly known as Alang-Alang, and 
Faek (Vetiveria sp.). 

Kha is extensively used by local people in every 
part of the country, whereas the use of Faek is 
limited to central and northeastern parts. The 
method of preparation is similar. After the 
grasses are reaped and dried in the sun, they are 
made up into bundles. Six to eight leaves are 
folded along one-third of their length on a piece 
of bamboo about one meter long and are secured 
by a string of Paw (Sterculia sp.). In certain 
parts, branches of Khon Tha (Harissonia per- 
forata) are used instead of bamboo. The thatch 
lasts for two to three years, and it is interesting 
to note that after the grasses are reaped the whole 
area is burned down, as it is thought that this 
will produce a good crop for the next season. 





Royal Forest Department, Bangkok, Thailand. 

Ancient Thai literature often described the 
forests as abounding in food plants and fruit 
trees. In the well-known story of Vesandorn 
Chadok, for instance, the part of the Himapharn 
forest into which Prince Vesandorn was exiled by 
his Royal Father, King Son Chai of Siplee, was 
described as full of fruit trees like wild banana, 
langsat (Lansium domcsticun i ) , Mafai (Baccau- 
rea sapida), Krathon (Sandoricwn koetjape), 
Lamyai (Euphoria longana) etc., on which 
the Prince could manage to survive without much 
trouble. It is felt that there is some truth in this, 
for, in the northeastern region of Thailand, 
especially, villagers more or less depend on the 
forest for their daily meals. Greens, bamboo 
shoots, mushrooms, etc., are gathered from the 
nearby woodland. They are cooked, roasted, or 
fried with either fish or meat and eaten with 
rice, the staple food of the teeming millions of 
the East. 

During his tours through the forests of Thai- 
land, the writer has always been interested in 
food plants, and, in order to give travellers and 
adventurers an idea of how one could survive 
when lost in the forest, he has compiled a list of 
plants from which food material can be obtained. 
The list is by no means exhaustive, and the com- 
mon herbs or fruit trees which are already well 
known and not actually growing wild have been 
omitted. The edible parts of the plants are either 
the flowers, fruits, shoots, leaves, stem, roots, 
bulbs, or tubers. 

For practical purposes, the forests of Thailand 
may be classified into four main types, namely, 
the evergreen forest, the mixed deciduous forest 
(including teak forest), the deciduous dipterocarps 
forest, and the mangrove forest. For the sake 
of convenience, the plant names listed here have 
been grouped under the various types of forests 
in which they are found. 

Table I. 
Food plant of the evergreen forest. 


Local name 

Bot. name 

Edible part 

How prepared 




Kum Nam 

Crataeva nurvala 

Young leaves 

Pickled in vinegar 

Almost taste- 

A medium-sized 

/ ' * \ 






Musa spp. 

"Stem" as a 

Taken fresh 

Wild banana is ra- 

^nmfii M^ 

source of water, 

ther full of seeds 

also fruit 



Garcinia cowa Roxb. 

Young shoots and 

By boiling with 

Slightly sour 

A medium-sized 

f \ 


meat or pork to 

tree in Southern 

VTJ/JJ 13) 

form an appetiz- 


ing broth 



Premna integrifolia 

Young shoots 

To be boiled or 

Crispy and al- 

A medium-sized 

y 4 


baked before be- 

most tasteless 


^ ' 

ing taken with 

chilli paste 



Arenga pinnata Merr. 

Young shoots 

May be taken 


A palm with simi- 


fresh or boiled 

lar appearance to 

nipa palm 



Amorphophallus spp. 

Young stems 

May either be 


A herb; must not 

( \ i f*^ 

fried or used as 

be washed with 

an ingredient in 

coldwater. The 

the preparation 

contact with cold 

of Thai curry 

water creates a reac- 

tion that causes a 

irritable itch in the 

throat when eaten. 

Only boiled water 

must be used. 

t Presented by T.P. Bank II. 



Bot. name 

Edible part 

How prepared , Taste 



Local name 



Nymphaea lotus L. 


May be taken ! Almost taste- 
fresh or fried or less 
boiled ' 

Aquatic; found in 
ponds or swamps 


Phak Koot 

Athyrium esculent um 

Young leaves 

To be boiled and 
prepared as an or- 
dinary vegetable 

Almost taste- 

A fern 



Baccaurea sapida Mu- 


Taken fresh 

Sweet or acid 

A forest fruit tree 



F/CWJ scandens Roxb. 


Taken fresh 


Sort of a fig tree 




Aganosma marginata 
G. Don 

Young shoots 
and leaves 

Taken fresh with 
chilli paste or 
pla-ra (fish paste) 

Slightly astrin- 

A scandcnt climber 


Wild Raspberry 


Rub us spp. 
Calamus spp. 

Young shoots 

Taken fresh 

To be boiled and 
taken with chilli 
paste or Pla-ra 
(sort of sal ted fish) 

Slightly bitter 

A shrub usually 
found in mountain 
evergreen forest 




Saraca indica Linn. 

Young leaves and 

To be used as an 
ingredient for pie- 
paring aThai curry 

Slightly sour 

Small tree growing 
along stream banks 


Hucha-niang or 


Pithecolobium jiringa 

Seeds both young 
and germinated 

Slightly astrin- 

Taken raw with 
chilli paste and 

Tree 15-20m tall 



Parkia javanica Merr. 

Germinated seeds 

Slightly astrin- 

Taken raw with 
chilli paste and 

Tree 30-40 m tall 



Parkia speciosa Hassk. 


Slightly astrin- 

Taken raw or 
pickled, with 
chilli paste and 




Elateriospermum tapos 

Mature seeds 

Slightly sweet 

Pickled and tak- 
en with curry 
or used in salad 

Tree 20-30 m tall, 

Table 2. 
Food plants of the mixed deciduous forest. 




Local name 

Bot. name 
Careya arborea Roxb. 

Edible part 

How prepared 


Kradone or pui 

Young leaves 

Taken fresh 

Slightly sour 

A medium-sized 





Sandoricum koetjape 

Dolichandrone crispa 


Taken fresh 

To be boiled or 
roasted before 
being taken with 
sauce or chilli paste 

Sour and slight- 
ly astringent 

Almost taste- 

A big forest tree 



Pi/w sp. 

Young shoots 

To be boiled be- 
fore being taken 
with sauce or 
chilli paste 


A vine 



Local name 

Bot. name 

Edible part 

How prepared 




Dillenia indica Linn. 

Young unripe 

Used as an ingre- 
dient in prepar- 
ing a Thai curry 


A large tree 


Cratoxylon polyan- 
thum Korth. 

Young leaves 

May be taken 

Slightly sour 

A medium- sized 

Bamboo shoots 
Bamboo Seed 

Dendrocalamus spp. 
Bambusa spp. & Oxy- 
tcnanthera spp. 


Seeds in the form 
of grains like rice 

Usually boiled be- 
fore being eaten 

Boiled like rice 

Almost taste- 

Like rice; may 
be substituted 
for rice in time 
of famine 

Found scattered 
about on the forest 
floor in bamboo 
flowering areas 


Dioscorca hispida 


Boiled with salt 

Starchy like 

May cause nasty 
itch in the throat 
if not properly 

Peka or Marid- 


Oroxylum indicum 

Seed in the young 

By boiling 

Almost taste- 

A shrub 





Spondias pinnata 

Tamarindus indica 

Young leaves and 

Young leaves and 

May be eaten 

May be eaten 

Slightly sour 
and astringent 


A big tall tree 

Found growing 
wild in old village 
sites; a big tree 

Lcb yciw 
(ifl 111)10 in) 

Zizyphus oenoplia 


May be eaten 


A thorny shrub 


Moringa oleifera 

Young pods and 

To be boiled be- 
fore eating; may 
be eaten fresh 

Almost taste- 

Found growing in 
old village sites 

with chilli paste 

Wan Poh 

Kaempferia galanga 


May be eaten 
fresh with chilli 

Crispy and al- 
most tasteless 

A herb 



(VI VI) 

Syzygium cumini 

Antelaea azadirachta 


Young leaves and 

Eaten fresh 
By boiling 

Rather bitter 

A big tree 

A medium sized 


Terminalia chebula 


Taken fresh 

Slightly acid 
and astringent 

A big tree 


Lagerstroemia macro- 

Young shoots 

To be boiled be- 
fore eating with 
chilli paste or pla- 
ra (fish paste) 

Almost taste- 



Piliostignaa malaha- 
sica Benth. 

Young leaves 

May be boiled 
with meat or pork 
to make a tasty 


A large shrub 


Acacia insuavis Lace 

Young shoots 
and inflorescence 

May be boiled or 
baked or taken 
fresh with chilli 

Slightly bitter 

Large wood and 
thorny, scandent 

paste or sauce 


Curcuma sp. 


Taken fresh or 
steamed with chil- 
li paste 

Slightly sweet 
and aromatic 

Herb with bright 
red bracts 




Table 3. 

Food plants of the deciduous dipterocarps forest. 


Local name 

Bot. name 

Edible part 

How prepared 




Kabok or Mameun 

Irvingia ma lay ana 

Endorsperm of 

By roasting until 

Like melon 

A big tree 



the seed 

well cooked 




Meliantha suavis 

Young leaves 

By boiling into 

Sweetish, de- 

A shrub. At times, 



a broth or used 


especially in the 

in a curry 

rainy season, it 

may turn deadly 

poisonous a 

mystery to both 

villagers and bo- 




Amorphophallus spp. 

Stems By boiling or 

Like lotus 

A herb with flow- 


frying with meat 
or pork (if avail- 

stem, almost 

ers in the form 
of spadises 





Costus speciosus 

Young shoots 

By boiling until 

Like asparagus 

A monocot herb 



well cooked 




Shorea talura Roxb. 


Used in a curry 

Slightly astrin- 

A big tall tree 




Table 4. 
Food plants of the mangrove forest. 


Local name 

Bot. name 

Edible part 

How prepared 




Kong-kang or 


Phizophora mucronat 

Young shoots 

By boiling and 
taking with sauce 
or chilli paste 

Just crispy and 
almost tasteless 


Kilek-pa or 
,* d ft * 


Cassia garrettiana 

Young leaves 
and flowers 

By boiling 

Slightly bitter 

A small tree 




Barringtonia asiatica 

Young shoots 

May be eaten 

Slightly astring- 

Found in swamps 


Samet Kao 

Melaleuca leucaden- 
dron Linn. 

Young shoots 

By boiling be- 
fore taking with 
chilli paste 


T.P. BANK ii : Would botanists and ethnobotanists 
study the ritual and folklore attached to these plants, and 

not leave it at a list of plant names? 





Department of Agriculture, Ministry of Agriculture, Bangkok, Thailand. 

Leaves of the palm known in Sanskrit as tala 
or tal were used in India as writing material for 
a very long time. The name tala or tal was con- 
fined to Borassus flahellifer, Corypha umbraculi- 
fera and Phoenix syhestris. When the Indians 
brought the custom of writing on the tala leaves 
to the Malesian area where Borassus flabellifer 
was plentiful, they also introduced that word 
into the local language. Tala became ron-tal 
and later, by metathesis, lontar; that is how the 
common name lontar, or lontar palm for Boras- 
sus flabellifer was derived. 

Whereas in Ceylon the leaves of Corypha wn- 
braculifera, the talipat (tali or tal, and pat = 
leaves), were used, because the Indian who 
brought the custom of writing on the tala leaves 
used the Corypha leaves, Corypha unibraculifera 
being plentiful in Ceylon. The word talipat 
later became talipot and the tree is called by the 
Anglo-Indian the talipot palm. 

The use of the talipot palm leaves as writing 
material was very likely adopted in Ceylon before 
B.E. 900, because it was known at that time that 
the manuscripts of all the teachings of the Lord 
Buddha had been written on talipot palm leaves. 

Thailand had for a long time been connected 
with Ceylon through religious channel. She 
accepted Buddhism from Ceylon as national 
religion. Some Thai monks were sent for training, 
and on return they were supposed to bring back 
authentic Buddhism to Thailand. It was recorded 
that religious manuscripts written on talipot palm 
leaves were brought into the Thai Kingdom just 
a little before B.E. 1800. Copies were then made 
using locally prepared talipot palm leaves vernacu- 
larly known as "Bi-larn". The Thai have used 
"Bi-larn" for writing religious manuscripts from 
that period up to the present. 

Talipot palm (Corypha umbraculifera) is found 
indigenously in Thailand. It forms great groves 
usually in the mixed, more or less dry and ever- 
green forests. It is a big tree of erect cylindrical 
trunk, 2-3 ft in diameter and 30-80 ft in height, 
clothed throughout with petiole bases. Leaves 
with stout petioles 5-10 ft long, lamina large 8-16 
ft in diameter palmately pinnatifid, plicate, cleft 
to about the middle into 80-100 linear lanceolate 


acute or bi-fid lobes. The lobe is web-liked blade 
in between two small ribs (veins). Flowering and 
fruiting occur after the age of 30 years. The plant 
dies after fruiting. 

Talipot palm is classified as one of the forest 
products which is a national treasure. Those 
who want to exploit this particular palm must 
apply to the Royal Forest Department for 
licenses. Under the rules and regulations of the 
Department, a license is valid for 3 years. After 
a permission is granted the exploiter may enter 
the Corypha grove and starts cutting juvenile 
plicate leaves with a curved and long handled 
knife. There arc 2 or 3 usable leaves in a tree, 
but 1 or 2 leaves only can be cut near the base of 
the lamina, leaving the youngest for future growth. 

All cut leaves are then recut at both ends. Only 
the middle part of the leaves which are about 
27 inches in length are gathered and dried in open 
air for 3 days, and then split along the ribs into 
bi-lobed slips (2 lobes with 1 rib in between). 
Bundles of one thousand bi-lobed slips tied up 
near both ends are prepared and shipped to 
Bangkok for sale. 

The Bi-larn makers purchase these raw bi- 
lobed slips, and grade them according to the 
width. There are four grades varying from 2.25 
inches to 1.75 inches wide. The undersized slips 
are not used as writing material but are sold as 
plaiting and weaving fibre for hats and other 

After grading, the thin ribs of the bi-lobed 
slips are removed, so that one raw bi-lobed slip 
produces 2 sheets. These sheets are again selected 
in order to eliminate poor specimens. 

Each sheet is cut to standard size and two 
holes, seven inches apart, used for binding purpose, 
are made along the centre line of the sheet. (The 
normal standard sizes of the sheets in inches are 
1.75 x 20.5, 2.0 x 20.5, 2.0 x 21.0, and 2.25 x 22.5.) 
Five hundred sheets are bound up into one packet 
with talipot ribs piercing through the holes for 
guiding. The packet is then placed in a wooden 
frame and tightly pressed in order to straighten 
and flatten the sheets. At the same time the edge 
are planed for smoothness. 

After that, the whole frames (with pressed 
packets of palm sheets) are placed in a smoking 
oven to be dried and coloured for about 48 hours. 
Rice husk or the palm leave waste may be used 
as fuel. The dried smoked sheets are then ready 
for writing. 

The so prepared talipot palm leaves are called 
"Bi-larn." In writing, this Bi-larn is placed on a 
metal table and characters are engraved with an 
iron stylus. Usually there are only 4 to 5 written 
lines on a single slip. To render the characters 
more legible, cow-dung or miller charcoal or soot 
mixed with coconut oil is smeared on the engraved 
leaf so that the written lines are blackened. 

The written Bi-larn are cleaned with sand and 
cloth and then bound into a packet by running 
through the holes two small cords, one for each 
hole, which are formed into two loops. The 
number of leaves in a packet depends on the size 
of the article or story. After binding into a book 
form, the edges are painted with a red dye-stuff 
known as "Chard", varnished with lacquer and 
then plated with gold leaves. These are very 


useful as a protection against fungi and insects 
which may occasionally attack Bi-larn. 

Nowadays, most of the religious articles or 
manuscripts are not transcribed, but printed. 
The use of iron stylus for writing on the palm 
leaf will be lost from the sight of younger gener- 
ation in the very near future. 

In printing, the prepared Bi-larn must first be 
roasted over a fire to remove excess oil in the 
leaf before it can be used by a printer. The 
characters are printed on both sides of the leaf, 
using generally 4 to 6 lines, and are arranged in 
three columns. 

The printed Bi-larn are brought together article 
by article and then bound into books. The edges 
are again dressed by planing, painting with red 
dye-stuff, and gold plated. 

The preparation of Bi-larn becomes a flour- 
ishing home industry in the Thai Kingdom; 
the palm leaves are used not only for religious 
and language writings, but also for making into 
cords, ropes, and plaiting or weaving material. 

Fig. 1. Talipot palm leaf. 



Fig, 2*' --Grading of the 


Fig. 3. Removing of the ribs from the bi-lobed slips. 



4 Cutting to 

Fig. 5. Trimming for uniformity. 



;;fr~ , 

V''.- j 2. ^ >j"' 

6, Binding in ihc 


Fig. 7. Planing for smoothness. 



In hot air 

Fig. 9. Removing oil over a fire before printing. 





Fig. 1 1 .Sorting the printed leaves for further binding. 




F.R. FOSBERO: I would like to ask Mr. Kasin about 
the geographic range of Talipot palm in Thailand. 

K. SUVATABANDHU: So far only in the North and 
Northeast, but in the South another species occurs, and 
all species are used for writing. 

C.G.G.J. VAN STEENIS: I am glad to have heard the 
detail on the talipot palm, that its use for writing has been 
imported to Thailand from Ceylon. I only know it from 
Bali where, however, the leaves used are derived from the 
lontar palm ( Borassus flahellifer ) which has been used for 
it since Hindu times and has been introduced by Hindu 
culture and wandered together with the invaders. The Bali 
lontars are mainly on religion, medicinal prescripts, history, 

art, etc. and thus the old ones contain much written 
information. They have been a great subject for study by 

F.R. FOSBLRG: I would ask Prof. Van Steenis if the 
famous Sumatran manuscripts, such as Prof. Bartlett has 
collected, are written on this sort of material. 

C.G.G.J. VAN STEENIS: As to papers in North Sumatra, 
I do not know particulars. The palms (Corypha, Borassus) 
are not growing in the mountains where the Batah live, 
but they occur on the sub-seasonal northern coastal 
plains of Atjeh (northernmost Sumatra). 

M.L. STEINER: In the Philippines palm leaves and 
bamboo strips are used. 





Chief, Section of Botany and Zoology, Forest Products Research Division, Royal Forest Department, Bangkok, Thailand. 

The Thai people are, by nature, migratory, as 
can be seen from their history. During their 
migrations, they learned much from nature 
around them. At one time, however, they did 
settle down, but hostile and aggressive neigh- 
bours forced them to move. Their migration 
over a period of thousands of years stretch from 
the Altai Mountains in Mongolia to the present 


In their history, the Thai people made much use 
of poison in both war and peace time. Most of 
the poisons used came from plants, and in spite 
of the impact of Western civilization upon the 
Thais, they are still used today. 

The poisons now in use are for poisoning arrow 
tips, insecticides, and fish-poisoning, as follows: 

Parts usec 


Local name 

Botanical name 






Khamin kru's 

Anamirta cocculus 





& Seeds 

& Seeds 



A ntiaris toxicaria 






Sakae dong 

Cocculus laurifolius 




Evergreen and 

mixed forests 


Entada phaseoloides 






Prik pa 

Ervatamia corymbosa 




. . 




Bua khru'ng sik 

Lobelia chinensis 



Evergreen for- 

est & cultivated 

Makham di khwai 

Sapindus rarak 





Tatum bok 

Sapium insigne 



- . 

Evergreen and 

mixed forests 

Rak pa 

Seme car pus curtisii 





Nawn tai yak 

Stemona tuherosa 



Evergreen and 


mixed forests 

Khika daeng 

Trichosanthes brae teat a 




Mixed forests 

Nong khru's 

Strophanthus scandens 






Croton tiglium 




Mixed forests 

or small 



Melanorrhoea usitata 



, . 

Mixed forests 


Melia azedarach 



Bark & 

Mixed forests 

& Seeds 


Salaeng chai 

Strychnos nuxivomica 




Mixed forests 

Wan nam 

Acorns calamus 





Albizzia procera 




Mixed forests 

Lai nam 

Derris elliptica 






Khao san 

Phyllanthus columnaris 







Local name 

Tatum thale 
Man kaew 

Botanical name 

1 Excoecaria agallocha 
Pachvrhizus erosm 


Parts used 

arrow j insect [ fish 
Tree j Latex i Latex 



Leaves Cultivated 
i & seeds 





Mae Malai Farm, Mae Rim, Chiang Mai, Thailand. 

The genus Camellia was originally thought to 
be indigenous to China, but later discovered that 
the location of Nature's original tea-garden is 
in the monsoon districts of Southeast Asia. The 
monsoon, one of the most vital producing factors 
to this part of the world, is called "Morasoom 
Tawan Tok Xieng Tai" or Southwest monsoon 
in Thailand, it blows across the Indian Ocean 
towards eastern Tibet, world's top-most plateau, 
carries moisture-laden clouds over the northern 
mountain ranges of Thailand, precipitates along 
its route as rains to an average of well over sixty 
inches per annum. 

The climate in Chiang Mai, Chiang Rai, 
Lampang, Mae Hongson, Nan and Phrac may be 
divided into three distinct seasonal categories; 
the cool and dry lasting from November to 
February, the hot and dry from March to early 
May, and the hot and humid from late May to 
October. The season in these vicinities is fa- 
vorably comparable to many tea producing 

The topography of northern Thailand may be 
divided into three regions: (1) The low-land or 
paddy field region, 'Tung Na." (2) The dipter- 
ocarpus region, "Pa Daeng." (3) The evergreen 
region, "Pa Dong Dib," with an altitude ranging 
from 200 to 300 metres MSL, 300 to 800 metres 
MSL, and 800 to 1,500 metres MSL respectively. 
The one that is concerned most in this report is 
the evergreen mountain region, the soil with a 
pH value of 4.5 to 5.5 of which has its origin 
from Gneisses rock and Quartzitic sandstone. 

Among these high hill ranges, there are tracks 
after tracks of Tea-trees growing. Villages after 
villages sprang up with the sites, people engaged 
in preparing the tea-leaf into "Miang," this name 
also implies to Shan, Laos, "Leppetso" to 
Burmese, and "Pickled-tea" to the English speak- 
ing world. 

"Miang" is consumed by Thai, Burmese, Lao, 
Shan, and Singpho (a tribe inhabiting the upper- 
reach of the Chindwin river in Northwest Bur- 

The miang trees have not at all been cultivated, 
they were allowed to grow in their natural states 
and some acquired a height of from 3 to 6 metres, 
when they are found too high for Plucking, the 


top branches may be broken off or given a light 

Tn preparing "miang" the leaf is plucked, in 
halves without stalk after the buds are all opened 
and not as a bud and two leaves, four times a 
year. The first plucking is locally called "Miang 
Hoa Pee," lasting from April to June, the second 
plucking "Miang Klang" from July to August, 
the third plucking "Miang Soi" from September 
to October and the fourth plucking "Miang Moey" 
(Dew Tea) lasting from November to December. 

After plucking and bundled, the leaves are 
put into a round wooden container "Hai Nueng" 
with plaited bamboo base and placed over a pan 
of boiling water for steaming which last from 10 
to 15 minutes. When the steaming process is 
through the content is thrown on to bamboo 
mat, people, sitting around kneading and retight- 
ening the bundles and let cool for 5 to 10 minutes; 
then the bundles or the day's plucking are placed 
and stamped down in bamboo silo 5 to 6 feet 
in diameter and 8 to 10 feet high. 

The silo is made of plaited bamboo lined 
heavily with wild plantain leaves, and kept adding 
to the content with steamed-leaf throughout the 
season ; at each filling the contents must be pressed 
down by tramping and planked with heavy stones. 
Under pressure, no doubt, the juice is oozed out, 
it is collected, boiled to a very thick consistency 
called "Nam miang" which is used as one of 
"Miang" delicacies. The leaf may be left in the 
silo for a period varying from 15 days to several 
months or until the commencement of the next 
season's plucking. 

Raw "miang" is very pungent and people pre- 
fers longer pickled one that has a sour taste. 
Miang has the smell of wet humus decomposed 
in a limited air supply (ensilage). It also has a 
slight sweet smell of fermented with yeast steamed 
glutinous rice (Khao mark) or alcoholic fermen- 
tation (zymurgy). The colour is bright yellowish 
green when taken out from the silo and rapidly 
becomes red when exposed for any length of 
time, the quality deteriorated since then. For 
preservation and handyness in packed-bull tran- 
sport from the producing hills down to the nearest 
motorized road or bullock-cart track, miang is 
repacked into bamboo baskets shaping like a 



mortar in lots of 60 to 80 kilograms. To assure 
of having proper insulation the basket is both sides 
smeared with viscid cow-dung, and heavily 
lined with plantain leaves. Once the basket is 
packed it never can be inspected in any transac- 
tion until retailing to consumers. Quality is 
guaranteed through honesty, the number of 
bundles in the basket is recorded by a numerical 
system engraved on a piece of bamboo stripe in 
the shape of a multiplication sign, a plus sign, 
a minus sign and dot sign ; each symbol represent- 
ing the amount is well known to all engaged in 
the trade. 

The marts of the pickled tea are in Mae Rim, 
Doi Saket, Ban Mae Ai in Chiang Mai district, 
Cheh Horn, Lampang district and Wieng Papao 
in Chiang Rai district. The storage for miang 
in these marts is a very interesting sight, a huge 
ditch swimming-pool-like is dug within the com- 
pound and kept filled with water, all tea baskets 
are kept submerged under water to keep them 
from contacting with the open atmosphere. 

Miang has a small export through Raheng- 
Mae Sot land route to South Burma, the rest of 

which is consumed locally. The common way of 
consuming "miang" is to wrap the leaf with 
salt and just hold in the mouth like lozenges, in a 
deluxe fashion "miang" is wrapped with hog's 
fat, pickled garlic, cocoa-nut-chips, ground-nuts 
and salt. The fashion of consuming miang is 
slowly dying out. 

Thailand is undoubtedly a country where tea 
could be grown in plantations commercially and 
economically. Hill-tribes labour is plentiful and 
who when employed to permanent agriculture will 
be a great asset in minimizing the destruction 
of forest wealth through "clearing culture" 
(Kaingining) or "Rai luen loi." 

In 1938 a small plantation was started the 
progress of which is rather slow. At Huey 
Tart a factory was built postwar time with modern 
equipment to make black tea with leaf purchased 
from smallholders. The production in 1952 was 
nearly 1 50,000 Ibs. The product is sold in Thailand 

Developments and researches can be carried 
on and on for further advancement. 





Australian National University, Canberra, A.C.T., Australia. 


The sugar palm (Bargot), Arenga pinnata 
(Wurmb) Merr., is found throughout Upper 
Mandating, Southern Tapanuli, Sumatra 1 . The 
area is one of complex mountain ranges bisected 
by a rift valley. Height varies from 400 to 2,000 
metres. The more temperate, wetter mountains 
contrast with the hotter, drier valley. The Medan- 
Padang road follows the latter, linking the main 
market centres and providing the import and 
export channel for the region. As the major 
subsistence crop, wet-rice in the major and tri- 
butary valleys, contrasts with dry-rice on the hill 
slopes. Rubber and coffee are the important 
cash crops. 

Table 1 shows the altitude and general soil 
type of the fifteen villages producing sugar for 

the extra-village market, throughout the year. 
The total number of villages in Upper Mandailing 
is eighty-seven. From the table it can be seen 
that economically significant palms grow on a 
clay soil usually of volcanic origin. The first 
nine villages listed are sited on lava flows ori- 
ginating from the volcano Sorik Merapi. It may 
be noted that most villages below the 500 metre 
level occur in valleys on soils derived from sandy 

The highest village in the area, Pagargunung, 
1,140 metres, is enclosed by hills rising to 1,310 
metres. Here the palm was previously utilised, 
but although the sap was sweeter than elsewhere, 
the flow is now considered too little to warrant 
collection. Here, too, coconuts were grown but 
did not bear. This situation is concomitant 
with Burkill's remarks that "the palm can be 

Table 1. 
Approximate height and general soil type of villages producing for the extra village market. 

Name of Village 

Si Banggor Djai 
Si Banggor Djulu 
Huta Tinggi 
Huta na Male 
Pasar Maga 
Angin Barat 
Si Antona 

Si Ladang Djulu 

Si Bio Bio 

Batahan Djulu 
Simpang Duhu Dolok 
Simpang Banjak Djulu 

Approximate height 
above S.L. in metres 






Note: Term used throughout the paper are those current in upper Mandailing. 

General Soil Type 
Heavy clay derived from lava 

Clay derived from limestone 
and schist 
from volcanic 
from volcanic 

,, from limestone 
,, from slate 

t Presented by C.S. Christian. 

1 Upper Mandailing includes the Ketjamatan Kotanopan and the three villages forming the Si Ladang complex at the foot 

of the mountain, Tor Sihite. 




sjown at greater elevations than the coconut..." 2 
During the month of fasting (Ramadan) in at 
east eight other villages there is a little produc- 
ion of sugar to supply the extra demand in that 
jeriod, and output is also increased in villages 
producing throughout the year. 

Formerly, nearly all villages in the area pro- 
iuced some sugar, but, with the introduction of 
'ubber, production has become concentrated in 
i few. The industry was stimulated during the 
Japanese occupation, when a ready market 
existed for sugar, but not for rubber. 

In recent years the planting of palms has become 
are. Distribution depends on wild animals, 
iuch as the civet cat, which eat the fruit and void 
he seeds. It is reported in many villages that 
he number of palms has increased in the past 
en years. The highest concentration of palms 
s found in the Si Ladang villages, the scattered 
elements of which lie within what is virtually one 
arge grove. Elsewhere in Upper Mandailing 
mly small groves occur and these outside the 
dllage proper. 


A palm comes into use when about ten years 
)ld. Before tapping commences, the common 
)eduncle of the male inflorescence (Sajatan) is 
>eaten with a wooden mallet ten times in a month, 
causing the flowers (Apili) to increase in size 
ind darken. This is taken as a sign that the 
ap (Aek Ni Bargot) is flowing into the inflores- 
cence and that it is ready for cutting. 

On cutting, one flower-spike (Arirang) is left. 
The heel of the knife blade is struck repeatedly 
vith a mallet to force it through the peduncle. 
Three days elapse between the first cutting and 
mnging a bamboo container (Taguk) below to 
collect the sap. Each morning a slice of the 
>eduncle about two cm thick is cut off. The 
low is then considerable, and a large container 
s set to catch the sap at night when the quantity 
exuded is large, while during the day a smaller 
container is used. 

The sap is collected every twelve hours. Each 
norning the peduncle is given two taps with the 
cnife handle before a slice is cut off, the size of 
vhich may be progressively increased to about 
hree cm. A leaf is tied round the cut to channel 
he sap into the container below. 

Access to a high peduncle is gained by climb- 
ng the palm or by means of a notched bamboo 

Substances may be placed in the container 
before it is fixed below the peduncle to retard 
fermentation, e.g., the fruit of the mangosteen 
(Manggis) or a pulverised root (probably ginger) 
from the forest. A further preventative measure 
is the washing out and smoking of all the bamboo 
containers before reuse. Clean containers are 
stored adjacent to the hearth. 

The length of time one inflorescence will yield 
is usually three months; but, where use is irregu- 
lar, it may extend to nine. The yield is measured 
in terms of the number of kilogrammes of 
sugar produced per day. Inflorescences in use 
(1956) were each yielding from one and a half to 
three kilogrammes of sugar per day. A very high 
price was being asked for an inflorescence yield- 
ing five kilogrammes. The female inflorescence 
(Alto) is distinguished, but never cut as the flow 
of sap from it is considered too insignificant. 


The fresh sap is boiled in large open pans in 
a special out-of-doors cooking place. After 
about one and a half hours, a thick liquid 
(Tangguli) is obtained which is returned to a 
bamboo container. This part of the preparation 
reduces the volume of liquid (one large pan of 
sap yields only one kilogramme of sugar) and 
prevents the sap from souring due to fermentation. 

When there is sufficient Tangguli on hand, a 
second boiling for one to one and a half hours, 
depending on the strength of the fire, produces 
a red liquid "free of water." To the boiling 
liquid a little coconut oil or crushed kernels of 
the candle nut (Aleurites moluccana Willd) or 
kernels from a bush (Djarat possibly Jatropha 
curcas L.) is added to prevent the liquid boiling 
over. A spoon with a perforated bowl may be 
used to remove froth in the early stages of cooking 
and any foreign materials. As the liquid thickens, 
a little is tested in the air on the stirring spoon 
to ascertain its setting properties and when ready 
it is ladled into moulds. 

The plank of wood on which the moulds are 
placed is washed down to prevent contamination 
from bacteria resulting from its prior use; this is 
seen in Mandailing eyes as a sickness of the sugar. 
The board may also be rubbed over with crushed 
candle nut kernels. The moulds (Bila) are hoops 
of bamboo made in three sizes, approximately 
7, 14, and 21 cm in diameter and 1, 2.5, and 3.5 
cm in height, respectively. 

I.H. Burkill: A Dictionary of the Economic Products of the Malay Peninsula, 1935, Vol. 1, p. 231. 



When set, the cakes (Paske) are pressed from 
the moulds and allowed to cool and harden. 
Cakes of like size are wrapped together in the 
dried fibre of the banana trunk and tied in parcels 
of a marketable weight. On account of the hy- 
groscopic nature of the sugar the parcels are 
stored above the hearth until marketed. Type and 
quantity of paske produced are adjusted to the 
market aimed at; the first, to the small scale 
intra-village spending of child and household; 
the second, to the market town and the largest to 
the requirements of bulk export. In 1956, the 
producers 1 selling price of a parcel of twenty 
medium sized cakes, was about Rp. 10, at about 
Rp. 2.50 a kilogramme. 

Variations in the process of cooking are mainly 
the result of differences in aptitude of the 
producers, in particular, in their ability to judge 
when the sugar has been sufficiently boiled. In 
the local markets the quality of the sugar varies 
considerably and, as a consequence, so does its 
price. Sugar may be marketed black in colour 
due to overcooking or smoke, moist and crum- 
bling from undcrcooking, or adulterated by vari- 
ous foreign bodies such as bees and palm fibre. 
Or, again, though carefully prepared, it may be 
held back until the market price rises, and then 
finally be sold in a deteriorated state. 

The site of the cooking place affects the quality 
of the final product. Cooking takes place in 
the precincts of the village, at the edge of the 
rice fields, or in the arboreal gardens. In the 
former case, unless sheltered by a roof and walls, 
dust may be carried by wind from the bare 
village streets into the cooking pans. In the 
garden and field sites it is usual to build a shelter 
to house the hearth and implements, but greater 
care has to be taken to prevent bees and other 
insects from being trapped in the liquid. But the 
construction of a cooking place in gardens is 
considered justified only where a number of palms 
owned or hired by the tapper are clustered 


The care of the palm and the preparation of 
the sugar are male occupations. Women assist 
indirectly by providing some of the wood for the 
cooking and directly in guarding the simmering 
liquid and tending the fire. Although women 
never participate in collecting the juice, in the 
Si Ladang villages women do climb the palms to 
collect the fibre. 



Ownership of the palms is established by 
inheritance, by planting, and by finding palms 
on land jointly owned by the village. Where a 
person desires usufruct of another's palm, several 
alternative arrangements exist. They are as 

1. The hiring of a tree and repayment with 
approximately a third of the final product. 

2. The buying of the usufruct of an inflores- 
cence at a fixed price after it has been estab- 
lished that the palm is yielding. 

3. The buying of the right to tap an untried 
palm. As there are many unproductive 
palms in the area, this represents a specula- 
tion. However, the slightly lower price is 
attractive, and it is said that some men have 
acquired the skill of being able to estimate 
the likelihood of a palm yielding. 


Palm sugar plays an insignificant part in the 
subsistence economy of the area. As a cash crop, 
its actual and potential use is severely limited by 
its storage properties which enforce a delicate 
adjustment between production and market 
demands. It has to compete with rubber and 
coffee for which a market is assured and which 
can be stored in anticipation of a favourable price. 
Hence the industry is usually combined with the 
production of other cash crops. In most of 
Upper Mandailing it is the preferred occupation 
of only a few and is irregularly practised. In the 
Si Ladang villages, on the other hand, tapping 
is regularly engaged in, and there is a greater 
standardisation in the quality of the product. 

In most areas the investment of labour and 
capital in the cultivation of the palm is slight as 
compared with that involved in coffee or rubber. 
Thus the palm is utilised when the demand is 
high, e.g., during the fasting month, and other- 
wise neglected. There is no loss due to idle 

Since the introduction of cheap white sugar 
with its better storage qualities, the use of this 
commodity has spread with a concomitant decline 
in the utilisation of the palm for sugar. 


The multipurpose functions of the palm are 
appreciated in Mandailing, all parts other than 


the roots being utilised, e.g., the fibre for roofing, food, the trunk for conduits, the fronds for 
cord, and brooms, the pith of trunk and seeds for shelters, and the leaves for decorative purposes. 

Note: Terms used throughout the paper are those current in Upper Mandailing. 


T. R. MCHALH: Is there any knowledge there concerning it was probably extracted from Arenga pinnata, not cane, 
the use of earthcrnware vessels for boiling sugar juices? If Did sugar techniques evolve from India or from Southeast 
sugar as a crystalline product was known before 600 A.D., Asia before Persian techniques were invented ? 





University of Michigan, Ann Arbor , Michigan^ U.S.A. 

In reviewing a vast amount of literature on 
primitive tropical agriculture, the writer has been 
impressed by the fact that there are a few chief 
types which must have been adapted to the early 
pre-agricultural environments of those tribes who 
became agriculturists. One finds himself spe- 
culating about where agriculture could have 
originated and what environments would have 
favored its development in succession to a purely 
food-gathering economy. It is a certain conclu- 
sion that primitive man in the humid tropics was 
not pastoral. The safest habitat for him would 
have been the shores of salt water, where he 
lived largely on molluscs and other marine pro- 
ducts, supplemented by what fruits and vegetables 
he gathered in the forest, as well as by hunting 
and by fishing. The first horticulture of the 
seaside dweller might have been the planting of 
such food-producing trees as coconut and bread- 
fruit; his first vegetable gardening the actual 
planting of edible plants that first presented the 
idea to him (or to her!) by springing up as weeds 
at the home site on rubbish heaps and other 
accidentally fertilized areas. Permanent or long- 
continued occupation of a site would result in 
constant accession of fertilizer derived from the 
debris of living, and so we might think of house- 
hold gardening and seaside horticulture as 
growing out of the sort of a subsistence that 
prevailed among people who built up shell 
mounds, inhabiting one place through many 
generations. It is easy to conceive of their 
migration along streams and into the forest as 
soon as they invented or came into possession 
of even simple tools and weapons for fishing, 
hunting, and defence. 

The fertility of burned-over .places would 
suggest the clearing of land, and moving the 
living site into the clearing would start an entirely 
new cultural evolution and population mobility, 
in which various forms of agriculture would 
coexist with the more ancient house-site gardening 
and horticulture. Early shifting dry-land (ladang) 
agriculture must have developed in areas of the 
most tropics where no irrigation was necessary, 
in forest clearings prepared by deadening trees 

without felling them and by the burning of slash. 
At first the crops must have been the same species 
that were gathered in the wild, such things as 
Tacca (in Polynesia) and yams (throughout the 
eastern tropics), and the tubers of edible aroids. 
A century ago heavy, edged, blade-like clubs not 
unlike canoe paddles were reported to be used in 
Fiji for slashing down the forest undergrowth in 
making agricultural clearings; and similar im- 
plements, still surviving elsewhere, indicate some- 
thing of the technique of clearing prior to the 
introduction of metal tools. Trees were deadened 
by ringing them with stone axes or adzes, or by 
piling and burning slash about them. The ashes 
of slash and the superficial organic debris of the 
forest floor fertilized the soil. Cultivation was 
abandoned when weeds and sprout growth 
became too rampant. There was no regular 
rotation of cropping with forest fallow. 

Ladang agriculture extended rapidly when the 
introduction of metal blades permitted the more 
complete clearing which is general in shifting 
agriculture today and which is characteristic of 
the Malay ladang. The greater area that could be 
cleared with the Malay metal-edged adze (beliung) 
or its equivalent enabled the chief men to have 
larger than ordinary clearings with more sub- 
stantial houses, built by the joint labor of the 
people. In Indonesia, the primitive clearing in 
which there would have been no tillage may have 
been called utna, and this term was retained in 
the original sense or as a name for the entire 
homestead, including clearing and house. 

The more gregarious people would retain one 
permanent house site, and around it the original 
ladang or uma would become stocked with fruit 
trees and little patches of disorderly mixed kitchen 
garden. Round about the old original inhabited 
site the repeated clearing of land for shifting 
cultivation might result in a regular orderly 
sequence of cropping and forest fallowing 
permanent land use of a sort. Over-population 
might result in the flocking away of a colony, 
to repeat the same process. In Sumatra such a 
colony would have been called a dusun. 

t Presented by T. P. Bank II. 



On the contrary, a less socially-minded, less 
gregarious population with plenty of land to 
spread over might never develop village life or 
orderly land use, but remain in a semi-nomadic 
state, utilizing previously unused forest as long as 
it lasted, and migrating as the old forest retreated. 
They would not have practiced tillage, but would 
merely have sown in the soft ash-covered soil. 

Chitemene and Rab agriculture grew out of 
typical shifting agriculture. In remotely separate 
places there has arisen the technique of increasing 
productivity of a clearing by burning upon it 
more slashings and other vegetable debris than it 
has produced during a period of forest fallow. 
Thus, in eastern tropical Africa a clearing is made, 
and in it are strewn branches cut from trees in 
the surrounding forest. With the debris thus 
supplemented, the ashes, sometimes dug into the 
soil with spades or hoes, are sufficient to produce 
good crops on sadly depleted soils. The technique 
is called chitemene. In India, an essentially similar 
form of supplementary fertilization by the ash of 
extra slashings from outside the cultivated area 
characterizes rah agriculture. 

The clearest transitions from ladang to per- 
manent dry-land agriculture by the introduction 
of tillage have been observed over the course of 
years in Java. Here the population has grown so 
enormously that the typical ladang has all but 
disappeared. Its place has been taken by gogo (or 
gaga) agriculture which is characterized by the 
introduction of tillage. Its counterpart in other 
parts of the world is generally designated as hoe 
agriculture or, in Africa, Bantu agriculture. 

It is unusual to find good descriptions of the 
primitive tillage of pre-plow agricultural peoples. 
Too often one runs across the expression "hoe 
agriculture" as indicating a stage of agriculture 
higher than the more primitive types of shifting 
cultivation, in which there is no tillage, but with 
no description whatever. The hoe agriculture of 
tropical Africa seems to have developed typically 
in forest lands marginal to savanna and more 
readily transformed by fire into grassland than 
likely to become regenerated forest if abandoned. 
So hoe agriculture has to have tillage and be able 
to combat grass as well as brush. It may have 
various developments before it develops into 
plow agriculture, one of which has developed 
both in India and Africa, namely sod-burning for 
securing ash from grass rhizomes and stubble, 
where wood is no longer available. The sods, 
dug by hoe or digging stick, are dried, then piled 
up and burned. There are enough rhizomes and 
roots so that the mass burns through, leaving a 

mixture of ashes and burned-out soil which is 
spread over the ground and dug in. 

So far all of the types of primitive agriculture 
considered may have evolved from a single point 
of origin. Two main groupings are on the one 
hand homestead or village horticulture with 
vegetable gardening which is more or less con- 
tinuous, and shifting agriculture, which may 
likewise give way to a type of permanent land use 
if population increases to the point where the use 
of land falls into a regular progressive rotation 
with alternate cropping and bush or forest fallow. 

Still left for consideration is wet-land rice 
growing, or sawah agriculture. What evolu- 
tionary sequence could have led to this? 

A chief objective of this communication is to 
point out the possibility that there may not have 
been any linear or connected sequence in the 
evolution of dry-land and wet-land types of 
agriculture which we may call the ladang and 
sawah types, respectively, using the convenient 
and rather familiar Malay terms, but rather, 
that wet-land rice agriculture may have had a 
direct origin from a food-gathering phase of 
human culture. Such a possibility is